Life history of Chnoospora implexa (Chnoosporaceae, Phaeophyceae) in culture

The life history of the brown alga Chnoospora implexa J. Agardh (Chnoosporaceae, Scytosiphonales) from Japan was studied in laboratory cultures. This species showed a heteromorphic and diphasic life history, alternating between erect gametophytes and discoid sporophytes. The gametophytes were dioecious and produced isogametes. The zygotes developed into sporophytes at 20°C under long‐day conditions, which formed plurilocular zoidangia. Zoids released from the plurilocular zoidangia developed again into sporophytes that always formed plurilocular zoidangia at 20°C and 25°C in long‐day conditions, and mainly unilocular zoidangia at 25°C in short‐day conditions. Zoids released from unilocular zoidangia developed into dioecious gametophytes. At 15°C zygotic erect thalli were formed and were revealed to be diploid by microspectrofluorometric measurements of nuclear DNA contents. The development and reproduction of unfused gametes were similar to those of zygotes. Some strains showed a direct‐type life history; gametophytic thalli were produced, but not via a sporophytic phase.     

LIFE HISTORY OF COLPOMENIA SINUOSA (SYCTOSIPHONACEAE,PHAEOPHYCEAE) IN THE AZORES1

Colpomenia sinuosa (Mertens ex Roth) Derbès and Solier (Scytosiphonaceae, Phaeophyceae) is a common species on the rocky intertidal shores of the Azores, where reproductive gametophytes occur throughout the year. Life‐history studies of this species were carried out in culture, and both sexual and asexual reproduction were observed. Anisogamous gametes fused to form zygotes. The zygotes gave rise to a filamentous prostrate sporophyte generation bearing unilocular sporangia, under both short‐day and long‐day conditions at 15 and 22°C, and to both unilocular and plurilocular sporangia, under the lower temperature condition. Unispores developed into gametophytes, and plurispores gave rise to filamentous sporophytes. Asexual reproduction was carried out by unfused female gametes and asexual plurispores produced from the same gametophyte. Unfused gametes developed into filamentous prostrate sporophytes producing unilocular sporangia in both culture conditions, and unispores released from the sporangia gave rise to gametophytes. Asexual plurispores from field gametophytes, under both culture conditions, developed directly into new gametophytes. The species exhibited three types of life history: a heteromorphic, diplohaplontic; a heteromorphic, monophasic (both with alternation between the erect and filamentous prostrate thalli); and a monomorphic, monophasic.     

DIFFERENTIAL LIFE HISTORY PHASE EXPRESSION IN TWO COEXISTING SPECIES OF SCYTOSIPHON (PHAEOPHYCEAE) IN NORTHERN CHILE1

The identity of two phaeophycean taxa that monopolized the middle‐lower rocky intertidal zone of a coastal area chronically exposed to copper mine wastes in northern Chile was unraveled. One of them was preliminarily identified as the gametophytic stage of Scytosiphon lomentaria (Lyngbye) Link. The other, a dark crust, resembled the alternate stage of some Scytosiphon species. Comparative analysis of morphology, life history, and DNA sequences strongly suggests that crusts corresponded to sporophytic S. tenellus Kogame and confirm that erect thalli belonged to S. lomentaria. A clear segregation of erect and crustose thalli was found using internal transcribed spacer region 1 and RUBISCO spacer sequences. Furthermore, whereas crusts always grouped with S. tenellus, erect thalli always grouped with S. lomentaria. Life history studies failed to connect the two entities. First, field‐collected S. tenellus produced progeny that either recycled the crust, which reproduced by unilocular zoidangia, or developed into erect thalli. The latter, unlike typical gametophytic S. lomentaria, developed patchy sori of plurilocular zoidangia. Second, S. lomentaria displayed a direct‐type life cycle, in which progeny from erect individuals only developed into erect thalli and produced only plurilocular zoidangia. This constitutes the first experimental study on Scytosiphon from the Pacific coast of South America and the first report of S. tenellus on this coast. It is also the first report of the crustose stage of Scytosiphon appearing as a perennial and dominant algal species in a temperate rocky intertidal system.     

A taxonomic study of Japanese Scytosiphon (Scytosiphonales, Phaeophyceae), including two new species

Morphological and culture studies were carried out on the genus Scytosiphon (Scytosiphonales, Phaeophyceae) growing in Japan. Morphologies of plurilocular zo-idangia and sporophytic thalli were found to be useful as taxonomic characters. Two types of plurilocular zo-idangial sori were recognized: (i) loosely coherent plurilocular zoidangia without a cuticular layer (loose type); (ii) tightly coherent plurilocular zoidangia with a cuticular layer (coherent type). The sporophytic thallus morphology was different among species in general appearance (tufted or discoid), degree of cohesion of erect filaments, and presence or absence of paraphyses. Two new species are described, Scytosiphon gracilis sp. nov. and Scytosiphon tenellus sp. nov, Scytosiphon gracilis is distinguished from other species by a thin medullary layer, coherent plurilocular zoidangia, no ascocysts associated with plurilocular zoidangia, and Compsonema-like sporophytic thalli. Scytosiphon tenellus is characterized by a thin medullary layer, coherent plurilocular zoidangia, ascocysts among plurilocular zoidangial sori, and Stragularia-like sporophytic thalli. Scytosiphon lo-mentaria (Lyngbye) Link is characterized by a constricted gametophytic thallus, loose plurilocular zoidangia, the presence of ascocysts, and sporophytic thaili identified with Microspongium gelatlnosum Reinke.     

Morphology and life history of Scytosiphon canaliculatus comb. nov. (Scytosiphonales,Phaeophyceae) from Japan*

Morphology of field material and life history in culture were studied in Scytosiphon canaliculatus (Setchell et Gardner) comb. nov. from northern Japan. Erect gametophytes of S. canaliculatus are cylindrical, tubular, up to 7 mm wide and 40 cm long, and without regular constrictions. S. canaliculatus has pronounced anisogamy and ascocysts accompanied with plurilocular gametangia. The life history of S. canaliculatus showed an alternation between erect gametophytes and crustose prostrate sporophytes bearing unilocular sporangia. Since field sporophytes of S. canaliculatus were found to be identical with Hapterophycus canaliculatus Setchell et Gardner (Ralfsiaceae, Phaeophyceae), it is proposed to transfer H. canaliculatus to the genus Scytosiphon. In the field, gametophytes with plurilocular gametangia appeared in spring and disappeared in summer. Sporophytes with unilocular sporangia were collected in late autumn and winter. Unilocular sporangia were produced at 15°C in short-day culture conditions and unispores developed into erect gametophytes at 5–15°C. It is suggested that the seasonal Occurrence of gametophytes in the field is due to the seasonal formation of unilocular sporangia, which is regulated by temperature and photoperiod.     

MORPHOLOGY AND LIFE HISTORY OF SCYTOSIPHON LOMENTARIA (SCYTOSIPHONACEAE,PHAEOPHYCEAE) FROM THE AZORES1

Morphological and culture studies of Scytosiphon lomentaria (Lyngbye) Link and Microspongium gelatinosum Reinke (Scytosiphonaceae, Phaeophyceae) were undertaken on material collected on the Island of São Miguel, Azores, where both species were commonly found. Erect thalli of S. lomentaria collected in the field were up to 33 cm long and 2.3 mm wide, tubular, hollow, and commonly constricted at intervals. The plurilocular sporangia were positioned in continuous sori on the thallus surface. Ascocysts were present. In the field, M. gelatinosum formed crustose to slightly pulvinate plants, were spongy in texture, and dark brown to black in color, which were circular or irregularly spreading over several centimeters and firmly attached to the substratum. Sessile unilocular sporangia were located in sori on the crust surface. In culture S. lomentaria plurispores developed into Microspongium‐like crustose prostrate thalli that formed unilocular sporangia. Unispores developed into new erect thalli that formed plurilocular sporangia. Sexual reproduction was not observed. In culture, M. gelatinosum unispores developed into erect thalli identical with S. lomentaria. These results are similar to those reported for other areas and suggest the occurrence in the Azorean plants of a monophasic and heteromorphic life history, involving both entities studied.     

Life history of Japanese Stypocaulon durum (Sphacelariales, Phaeophyceae)

Phenology, morphology, life history and responses to different temperature and photoperiod conditions were studied in Japanese Stypocaulon durum (Ruprecht) Okamura. Erect thalli of the species were collected year-round, but the mature thalli forming either uniloc-ular sporangia or two different types of plurilocular structures (evidently gametangia) on separate thalli were found only in winter. ln culture, an isomorphic life history is suggested for the species, alternating between a sporophyte forming unilocular sporangia and gam-etophytes forming plurilocular macro- (female) and micro- (male) gametangia. Contents of unilocular sporangia were not released, but germinated in situ, developing into erect thalli forming plurilocular gametangia. Macrogametangia released aplanogametes (oospores), but male gametangia appeared to be non-functional, although flagellated cells were once formed in the loc-uli. This is the first report of plurilocular gametangia in the species. Although the species grew well and matured under considerably lower temperature conditions than European Stypocaulon scoparium (L.) Sauvageau, its temperature requirements showed similarity to northwestern Atlantic Stypocaulon species. This supports the notion that northwestern Atlantic Stypocaulon is conspecific with S. durum.     

First record of Asteronema rhodochortonoides (Phaeophyceae) for the Pacific Ocean

Morphological observations of a minute, filamentous, branched brown alga epiphytic on Sargassum thun‐bergii (Mertens ex Roth) Kuntze were made on material collected at Tsuyazaki (33°48′N, 130°27′E), Fukuoka Prefecture, southern Japan. This alga was assignable to Asteronema rhodochortonoides (Børgesen) Möller et Parodi in having stellately arranged chloroplasts with several pyrenoids grouped in the center, predominantly apical growth, narrow filaments, and elliptical or broadly elliptical plurilocular zoidangia that are apically or laterally formed on upright filaments. A comparison of partial nuclear small subunit rDNA sequences between the Japanese material and A. rhodochortonoides from the Canary Islands showed only two or three nucleotide differences. This supports our assignment of the Japanese material to this species as a first report for the Pacific Ocean. In laboratory cultures, zoids released from plurilocular zoidangia developed into plants with morphology similar to the field‐collected plants. This cycle repeated without production of unilocular zoidangia in our cultures.     

Morphology and life history of Halopteris filicina (Sphacelariales, Phaeophyceae) from Korea

The vegetative morphology and life history of Halopteris filicina (Grateloup) Kutzing, collected from Korea, were examined in laboratory culture. Field plants attaining 3–5 cm in height were epilithic, tufted, yellowish-brown, and produced numerous erect axes with alternately distichous branches from compact basal discs. They were cultured under a 12:12 h LD photoperiod at 10°-C, 15°C and 20°C to observe the influence of temperature on reproduction. At 10°C plants grew only vegetatively, whereas at 15°C and 20°C they produced unilocular sporangia. Unispores released from sporangia developed into monoecious, anisogamous gametophytes that formed plurilocular female and male gametangia on the same lateral branches. The zygotes, by fusion of female macrogametes and male microgametes, developed into sporophytes bearing unilocular sporangia, whereas the unfused female gametes germinated parthenogenetically. This species was confirmed to have an isomorphic life history, basically similar to the other species of Sphacelariales.     

Cladosiphon umezakii sp. nov. (Ectocarpales, Phaeophyceae) from Japan

The new species Cladosiphon umezakii Ajisaka (Ectocarpales, Phaeophyceae) is described from Japan based on morphology and DNA sequences. The species resembles Cladosiphon okamuranus Tokida in its gross morphology; somewhat slimy, cylindrical, multiaxial and sympodial erect thallus, arising from a small disc‐shaped holdfast, and branching once to twice. However, C. umezakii has considerably longer assimilatory filaments (up to 840 μm long, composed of up to 90 cells) than any known taxa of the genus. The species is a winter to spring annual, growing on lower intertidal to subtidal rocks of more or less exposed sites on the north‐eastern coast of Kyushu and on both the Pacific and the Sea of Japan coasts of Honshu. Specimens from the Sea of Japan coast had both unilocular and plurilocular zoidangia, whereas those from Kyushu and from the Pacific had only unilocular zoidangia. Unilocular zoidangia were formed on the basal part of assimilatory filaments, and plurilocular ones were transformed from the distal part of assimilatory filaments. DNA sequences of the Rubisco‐spacer (rbc‐spacer) region and the nuclear rDNA ITS region (ITS1, 5.8S and ITS2) supported the distinctness of the species.     

MOLECULAR PHYLOGENY OF DISCOSPORANGIUM MESARTHROCARPUM (PHAEOPHYCEAE) WITH A REINSTATEMENT OF THE ORDER DISCOSPORANGIALES1

A molecular phylogenetic analysis of the little‐studied filamentous brown alga Discosporangium mesarthrocarpum (Meneghini) Hauck using rbcL and partial 18S rDNA sequences revealed that the species forms a monophyletic clade with Choristocarpus tenellus (Kütz.) Zanardini that is sister to all other brown algae. Although D. mesarthrocarpum has unique disk‐shaped plurilocular reproductive organs, D. mesarthrocarpum and C. tenellus share the following basic morphological features, which are considered to be plesiomorphic characters in the brown algae: (1) apical (and diffuse) growth; (2) uniseriate, subdichotomously branched filaments; (3) multiple chloroplasts per cell without pyrenoids; and (4) lack of heterotrichy and of phaeophycean hairs. The rbcL DNA sequence of an Australian D. mesarthrocarpum specimen showed considerable deviation from Mediterranean and Macaronesian specimens. Therefore, the presence of a second species in the genus is suggested; however, the taxonomic treatment of this putative species is not pursued in the present report. Regarding the higher‐ranking systematic position of D. mesarthrocarpum, reinstatement of Discosporangiaceae and Discosporangiales is proposed, and the inclusion of Choristocarpaceae in the order is also suggested. Under short‐day and long‐day culture conditions at 15°C–25°C, Mediterranean D. mesarthrocarpum exhibited a direct type of life history, with a succession of uniseriate filamentous thalli bearing characteristic disk‐shaped plurilocular zoidangia, but thalli did not survive at 10°C and below.     

Morphology and life history of four Japanese species of Elachista (Elachistaceae, Phaeophyceae), including a new record of E. fucicola

Elachista fucicola (Velley) Areschoug (Elachistaceae, Phaeophyceae) is newly recorded from Japan, and compared with three previously described species, Elachista coccophorae Takamatsu, Elachista mollis Takamatsu and Elachista okamurae Yoshida. All species showed direct‐type life histories in culture without sexual fusion. Prostrate filaments of E. fucicola formed globular plurilocular zoidangia similar to those reported from Atlantic isolates of this species. However, these were different from uniseriate plurilocular zoidangia of the three Japanese species. Furthermore, the position of a meristematic region in assimilatory filaments and the morphology of paraphyses can easily distinguish these from E. fucicola. Previously emphasized differences in cell length : width ratios in assimilatory filaments showed only slight differences. Elachista coccophorae is characterized by thick‐walled assimilatory filaments and curved paraphyses, and the absence of downwardly growing rhizoidal filaments. Elachista mollis and E. okamurae are very similar. However, germlings from plurizoids formed on erect filaments of E. mollis showed characteristic pseudodiscoid growth in culture. Halothrix coccophorae Ohta and Elachista zosterae Noda are reduced to synonyms of E. coccophorae and E. mollis, respectively.     

The influence of light quality on the development of the brown algae Petalonia and Scytosiphon

Petalonia fascia (O. F. Müll.) Kuntze, P. zosterifolia (Reinke) Kuntze and Scytosiphon lomentaria (Lyngb.) J. Ag. have been cultured in white, blue and red light of equal quantum irradiance at 15°C. Hairs, knot-filaments and Ralfsia-like crusts were formed only in blue light, whereas the prostrate system of red-grown plants consisted entirely of sparingly-branched and mostly uniseriate filaments. The production of erect thalli from the prostrate system was controlled or stimulated by red light, but these erect thalli became fertile only in the presence of blue light. All three species exhibited all of these types of response, although specific differences in the degree of certain types of response were observed.     

79 Systematic revision of Phaeostrophion Irregulare (phaeophyceae) and proposal of a new family phaeostrophionaceae

Phaeostrophion irregulare S. & G. is distributed along the Pacific coast of NW America, and has been placed in the family Punctariaceae, order Dictyosiphonales (or Ectocarpales s.l.), based on morphological resemblances. Culture studies have indicated a direct life history, with erect thalli forming unilocular and/or plurilocular sporangia. However, the occurrence of a perennial prostrate system (holdfast), presence on the holdfast of marginal meristematic cells, and lack of pyrenoids in the chloroplasts, suggest that this taxonomic assignment is questionable. Our TEM observations confirm the lack of pyrenoids in the chloroplasts. The occurrence of reductive divisions in the unilocular sporangia was confirmed by comparing the DNA content in each nucleus using a fluorescent stain, but both unilocular and plurilocular sporangia-bearing erect thalli showed diploidy. Molecular phylogenetic analysis using rbcL (chloroplast) and 18S rDNA (nuclear) gene sequences reveals that Phaeostrophion is not included in the clade of Ecotocarpales s.l., but diverged relatively early in the evolution of Phaeophyceae, forming a clade with Sphacelariales and Syringodermatales, which share the apical/marginal meristematic mode of growth. Although we propose a new family Phaeostrophionaceae to accommodate Phaeostrophion , we suspend judgment of the appropriate taxonomic treatment at the ordinal level.     

Morphology and life history of Coelocladia arctica (Dictyosiphonales,Phaeophyceae), new to Japan*

The occurrence of Coelocladia arctica Rosenvinge (Dictyosiphonales, Phaeophyceae) is reported for the first time in the Pacific Ocean, from Oshoro, Hokkaido, Japan. The field material was approximately 30 mm in height, up to 300 M-μn in diameter, 2-3 times irregularly branched and provided with phaeophycean hairs and plurilocular sporangia. The thallus was composed of several (usually four) large, hyaline, rounded, isodiametric inner cells, with smaller sub-cortical and pigmented cortical cells. The plurilocular sporangia were 3-4 celled, often branched, and protruded from the cortical cells, arranged in a crown-shaped complex. In culture, the plurispores germinated unipolarly leaving an emptied original spore wall, and developed into a branched protonema. Cells of the protonema as well as the erect thallus contained several disc-shaped chloroplasts with pyrenoids. Uniseriate erect filaments arose from the protonema, then became polystichous and formed branches. Unilocular sporangia were never observed in the field or in cultured material. Erect thalli were formed under culture conditions of 5-15°C, and developed a thick parenchyma at 5-10°C, irrespective of the day length.     

Reproductive morphology and DNA sequences of the brown alga Platysiphon verticillatus support the new combination Platysiphon glacialis

Platysiphon verticillatus, a brown alga endemic to the Arctic, was described based on vegetative specimens collected at Inglefield Bay, West Greenland. The species is distinctive in having a lanceolate blade‐like thallus terminated by a terete portion, both covered with hair‐like assimilatory filaments. Punctaria glacialis was described from Eastern Greenland, and the species differs from other Punctaria species in lacking hairs and plurilocular zoidangia. Unilocular zoidangia were reported, but instead of zoids being released they formed cell walls in situ developing the appearance of plurilocular zoidangia. However, the fate of the zoids, as well as the walled cells was not traced, and the life history of the alga has remained unclear. By comparing DNA sequences (cox1, cox3, and rDNA ITS2) of specimens morphologically referable to Platysiphon verticillatus and Punctaria glacialis collected at Baffin Island, as well as re‐examining morphology and studying crude cultures, we concluded that they are the same taxonomic entity. Furthermore, their cox3 sequence and vegetative morphology agreed with those of the type specimen of Punctaria glacialis. Consequently, we propose Platysiphon glacialis comb. nov. The life cycle could not be completed in culture, but we hypothesize that in situ germination of the unizoids produces reduced gametophytes housed in peripheral tissue of erect sporophytic thalli.     

Life history and systematic position of the rare brown alga Climacosorus mediterraneus

Cultures of Climacosorus mediterraneus were established with material from Spitsbergen. This species has previously been reported from three widely separated geographic areas: the west Mediterranean, north-west France, and north Norway. Although adapted to temperatures near 0°C, the northern isolate grew well at 15°C which approximates the summer temperature of the southern habitats. The life history proved to be entirely asexual with successive generations bearing unilocular sporangia on the erect filaments and plurilocular sporangia infrequently on the prostrate system. Swarmers of the two reproductive organs gave rise to similar progeny. Chromosome counts revealed equal numbers in somatic cells and unilocular sporangia which substantiates the development observed. The absence of true hairs and the few longitudinal divisions, which only occur in connection with formation of unilocular sporangia and branches, suggest that C. mediterraneus should be assigned to the Pilayellaceae.     

OBSERVATIONS ON THE LIFE HISTORY OF PAPENFUSSIELLA CALLITRICHA (PHAEOPHYCEAE,CHORDARIALES) IN CULTURE1

Papenfussiella callitricha (Rosenv.) Kylin from eastern Canada was studied in culture. Zoids from unilocular sporangia develop into microscopic, filamentous, dioecious gametophytes which produce isogametes in filament cells and few-chambered plurilocular gametangia. Unfused gametes germinate to reproduce the gametophytes. Fusion takes place between a settled (“female”) and a motile (“male”) gamete. The zygote gives rise to a filamentous plethysmothallus that reproduces asexually by zoids formed in thallus cells and in few-chambered plurilocular zoidangia. Erect macrothalli are produced on the plethysmothallus, beginning with the formation of upright filaments. Later on, these filaments become the terminal assimilators of the macrothalli. Further assimilatory filaments, rhizoids, and unilocular sporangia are produced in a branching region at the base of the terminal assimilator. Zoids from unilocular sporangia formed in culture germinate to reestablish the gametophyte phase. Chromosome counts yielded n = 19 ± 3 for the gametophytes, and 32 ± 6 for the sporophyte, both plethysmothallus and macrothallus.     

Polysiphonia perforans sp. nova (Ceramiales,Rhodophyta) from the Mediterranean Sea

ABSTRACT

A new species of the genus Polysiphonia from the Mediterranean Sea is described. It is an ecorticate species, with 4 pericentral cells, showing prostrate axes from which erect axes arise. Rhizoids are formed by pericentral cells in a median position remaining in open connection with them. Erect axes are straight, simple throughout or pseudodichotomous at the base then simple or with one (rarely two-three) orders of branching; trichoblasts and scar cells not observed. Only tetrasporangial plants were found. They show ellipsoid tetrasporangia borne in short straight series, often interrupted by sterile segments. The new species is also characterized by a peculiar habit consisting of prostrate axes, adhering to lower faces of Peyssonnelia spp. by means of rhizoids growing upward, from which erect upright axes perforating thalli of the supporting species arise. A comparison with the related species of Polysiphonia was also carried out.     

Morphology, cytology and taxonomic remarks of four species of Stigeoclonium (Chaetophorales, Chlorophyceae) from Argentina

Four species of Stigeoclonium from Argentina were studied by means of transmission electron microscopy and light microscopy. For species identification, we collected data related to the prostrate system and zoospore germination. We also determined the chromosome number for each species. Stigeoclonium aestivale showed a more developed erect system than the prostrate one, zoospore germination was predominantly of erect type and the chromosome number was 8. Stigeoclonium tenue presented well developed, erect and prostrate systems, zoospore germination was initially of the prostrate type and the chromosome number was 5. In Stigeoclonium variabile the prostrate system predominated over the erect one, zoospore germination was strictly of the prostrate type and the chromosome number was 3. Stigeoclonium farctum presented a more developed prostrate system than the erect one, zoospore germination was strictly of the prostrate type and the chromosome number was 8. The ontogeny of the zoospore germination was related to the final relative development of the prostrate and erect portions of adult thalli.