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.     

SPORANGIA OF RALFSIA FUNGIFORMIS (GUNN.) SETCHELL ET GARDNER12

Unilocular and plurilocular sporangia of Ralfsia fungiformis are, for the first time, recorded for North America. The validity of certain types of structures, considered by some authors as plurilocular sporangia, is discussed.     

Neoleptonema yongpilii E.-Y. Lee & I.K. Lee,gen. et sp. nov. (Phaeophyceae), based on morphological characters and RuBisCO spacer sequences

An elachistacean epiphyte, Neoleptonema yongpilii E.-Y. Lee & I.K. Lee, gen. et sp. nov., is reported from Korean coasts. The plants are distinguished by having unbranched assimilatory filaments with intercalary plurilocular sporangia as well as lateral plurilocular sporangia from the cortex. The new genus differs from the genera Leptonematella P. Silva and Halothrix Reinke by having pod-shaped plurilocular sporangia on the medulla, and from Elachista Duby and Proselachista Y.P. Lee & Garbary by having intercalary plurilocular sporangia and a poorly developed medulla. The phylogenetic relationships of Neoleptonema yongpilii were inferred from the spacer sequences between the genes coding for the large and small subunits of the RuBisCO gene. The new genus is a member of a poorly resolved clade consisting of several genera within the Elachistaceae, within which it is more closely related to Halothrix and Elachista nipponica than to Leptonematella.     

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.     

Life histories of Colpomenia sinuosa and Hydroclathrus clathratus (Scytosiphonaceae, Phaeophyceae) in culture

Life-history studies in culture were carried out on Colpomenia sinuosa (Mertens ex Roth) Derbès et Solier and Hydroclathrus clathratus (C. Agardh) Howe (Scytosiphonales, Phaeophyceae) from Japan. These species showed a heteromorphic life-history pattern with an alternation between erect thalli bearing plurilocular zoidangia and prostrate thalli bearing ectocarpoid plurilocular and unilocular zoidangia. Plurizoids released from erect and prostrate thalli developed into prostrate thalli. Unizoids, however, developed into erect thalli. Prostrate thalli produced plurilocular zoidangia in long-day conditions and unilocular zoidangia in short-day conditions at 10-20°C. Prostrate thalli of C. sinuosa formed ascocysts. Germlings of both species did not grow at 5°C.     

RELATION OF TEMPERATURE TO DEVELOPMENT OF THE MACROTHALLUS OF DESMOTRICHUM UNDULATUM1,2

Desmotrichum undulatum (J. Agardh) Reinke (Phaeophyta) was found in the winter months in the littoral zone of a tidal marsh in Virginia. When field collections were maintained in culture media at 21 C, most cells of the thallus developed into plurilocular sporangia. Released zoospores developed into microthalli from which clonal cultures were established. At 21 C microthalli produced sporangia which released zoospores, subsequently developing into more microthalli. However, microthalli at 6 C formed filaments which subsequently developed into thalloid plants. When the thalloid plants of Desmotrichum formed in culture were subjected to 21 C, plurilocular sporangia were formed. These results support the proposition that the seasonal periodicity of Desmotrichum is due to a temperature-regulated phenomenon in the development of macrothalli.     

The ship-fouling algaEctocarpus

Summary The unilocular reproductive stages of the brown algaEctocarpus have been examined with the electron microscope. One observation in particular, which is believed to have been made for the first time in Phaeophycean studies, is that the zoospore derived from unilocular sporangia possesses a microanatomy considerably different from that of spores formed by plurilocular sporangia. It is suggested that this difference may have an evolutionary significance and may lead to a better understanding of the life cycle ofEctocarpus.     

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.     

Sexual reproduction in British Ectocarpus Siliculosus (Phaeophyta)

One male and one female gametophyte of Ectocarpus siliculosus have been detected in a sample of 19 plants taken from the Isle of Man. Fertilization takes place between gametes formed in plurilocular gametangia. Clear cross-fertility exists with Mediterranean strains. This result confirms the notion that the concept of the gametic character of meiospores in Ectocarpus should be abandoned.     

CULTURE STUDIES ON REPRODUCTION OF SPERMATOCHNUS PARADOXUS (PHAEOPHYCEAE,CHORDARIALES)1

The life history of Spermatochnus paradoxus (Roth) Kütz. isolated from the Mediterranean Sea was studied in culture. Meiospores develop to a microscopic stage (microthallus) which at 20°C perpetuates asexually by plurilocular sporangia and formation of new microthalli. At 9°C microthalli act as homothallic gametophytes. Fusion of isogametes results in a diploid microthallus which, after differentiation of an apical cell, leads back to Spermatochnus plants. In addition, gametes develop without fusion to form haploid macrothalli, the further fate of which has not been determined. Chromosome numbers alternate between n = 20 ± 2 in the microthalli and 2n = 41 ± 4 in macrothalli.     

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.     

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 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.     

STICTYOSIPHON SORIFERUS (PHAEOPHYTA,DICTYOSIPHONALES) FROM EASTERN NORTH AMERICA1

Although earlier North American records of Stictyosiphon soriferus (Reinke) Rosenvinge cannot he substantiated, the species is shown to be common in Newfoundland and is also reported from Nova Scotia. Plants are summer annuals and a component of ephemeral subtidal communities characteristic of suitable inlets and harbors. Only plurilocular reproductive organs are produced. In culture the life history of populations from both sides of the North Atlantic consists of a direct replication by asexual zoospores. A microscopic stage might occur in the field, and is presently known only from European cultures. In eastern Canada S. soriferus is readily distinguishable from S. tortilis (Rupr.) Reinke, S. subsimplex Holden and S. griffithsianus (Le Jol.) Holmes & Batters.     

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.     

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.     

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.     

Extinction of nanicellid foraminifera during the Frasnian–Famennian biotic crisis: some far‐reaching evolutionary consequences

This study evaluates the severity of the poorly known and mostly underestimated foraminiferal extinction during the Frasnian–Famennian biotic crisis and its evolutionary aftermath. During this global event, worldwide, truly plurilocular planispiral (Nanicellidae) and uniseriate, palmate (Semitextulariidae) foraminifera associated with metazoan reefs died out entirely. Highly advanced test morphology such as that of nanicellids did not reappear in the earth's history until the Late Triassic. Moreover, morphotype comparable to that of the Devonian bilaterally flattened and palmate semitextularids appeared again until the Middle Jurassic (Frondicularia, Lagenida). In terms of the degree of test septation and chamber arrangement as well as general test shape, these foraminifera were ‘very far ahead of their time’. In consequence, foraminifera suffered a significant collapse during the F‐F biodiversity crisis, leading to an amazingly long evolutionary time lag in the case of plurilocular foraminifera lasting at least 150 million years.     

Studies on Kuckuckia spinosa (Fucophyceae, Sorocarpaceae): life history, temperature gradient experiments, and synonymy

Kuckuckia spinosa and Kuckuckia kylinii are distinguished mainly by the presence of basally sheathed true hairs in the latter. However, culture studies have shown that the sheath is an ephemeral structure as it splits, breaks off, and the basal remnant disintegrates and disappears eventually. Therefore, Kuckuckia kylinii gradually develops into Kuckuckia spinosa , also with regard to filament diameter. Consequently, the two species are merged with priority to K. spinosa. The life history is of the direct type as the swarmers from the plurilocular sporangia develop by immediate differentiation into plants similar to the mother plant. It has been confirmed experimentally that this species belongs to the warm temperate Mediterranean-Atlantic group with a southern reproductive boundary close to 25°C and a northern reproductive boundary close to the 10°C summer isotherms. This is the first report of this species from the Canary Islands.