Ecological studies on the community of drifting seaweeds in the south-eastern coastal waters of Izu Peninsula, central Japan. II: Seasonal changes in plants showing maximum stipe length in drifting seaweed communities

The authors examined seasonal patterns in 20 sargas‐saceous species (maximum stipe length of ≥45 cm) from 902 patches of drifting seaweeds in the southeastern coastal waters of the Izu Peninsula, central Japan, between spring and autumn, 1991–1993. The first analysis, dealing with plants occurring for three consecutive years, examined the top three ranked species determined from monthly pooled means of frequency of appearance in these 3 years. The second analysis examined not only the top‐ranked species in monthly means in each year, but also the continuity of appearance of species among years. The third analysis was made from dendrogram analysis based on monthly samples from each year. The first and second analyses revealed three seasonally discernible changes. From April to June, dominance shifted from Sargassum horneri (Turner) C. Agardh to Sargassum yamamotoi Yoshida with a stable transition; in July, S. yamamotoi, Sargassum macrocarpum C. Agardh, Sargassum crispi‐folium Yamada and Sargassum piluliferum (Turner) C. Agardh appeared consistently in an unstable domination with low frequency of appearance by Sargassum nipponicum Yendo, Sargassum fulvellum (Turner) C. Agardh, S. macrocarpum or S. crispifolium; in August to October, there was a stable shift from S. macrocarpum to Sargassum micracanthum (Kützing) Endlicher and Sargassum ringgoldianum Harvey. In the third analysis, monthly samples from each year were divided into four groups, characterized using the top five ranks. This characterization indicated that the top‐ranked species shifted from S. horneri to S. yamamotoi, to S. macrocarpum and to S. micracanthum with the change of seasons. Because of an overlap in the characteristic species in these three analyses, seven species (S. horneri, S. micracanthum, S. yamamotoi, S. macrocarpum, S. ringgoldianum, S. nipponicum and S. crispifolium) were regarded as dominant species. Results are compared to our previous study and it is suggested that the similarity in the pattern of plant community during seasonal changes over the period from April to June and in July resulted from both stable and unstable changes, respectively, but discrepancies in the pattern of plant community during seasonal change from August to October were the result of differences in the frequency of appearance of S. ringgoldianum and S. macrocarpum in August 1991. These results indicated that the seasonal changes of dominant species in the plant community from April to October in this study area were periodic except in July.     

Ecological studies on the community of drifting seaweeds in the south-eastern coastal waters of Izu Peninsula, central Japan. I: Seasonal changes of plants in species composition, appearance, number of species and size

Seasonal patterns of drifting seaweeds in the southeastern coastal waters of Izu Peninsula of central Japan were examined by sampling 966 patches from spring to autumn 1991–1993. In total, 57 plant species appeared, including 10 epiphytic algal species. Monthly totals of the number of species, excluding epiphytic aigae, were highest in May (33) and August (27), though 19–21 species of sargassaceous algae were found from May to August, The number of species, excluding epiphytic algae, in one patch of drifting seaweeds was 1 to 11 (x?= 2.93 ± 2.06) with high richness in May a result of almost entirely sargassaceous species. The wet weight of each patch and maximum stipe length of plants varied from 5 to 6970 g and from 20 to 840 cm (x?= 536.1 ± 782,3 g and 110.6 ± 76.8 cm), respectively, with highs in April and May. Out of 18 species common to all years, 10 species dominated the top or second rank in monthly pooled frequency of appearance. Seasonal changes of these 10 major species were examined, Sargassum horneri (Turner) C. Agardh and Hizikia fusiformis (Harvey) Okamura were abundant in April, but were replaced partly by Sargassum muticum (Yendo) Fensholt in May and largely by Sargassum yamamotoi Yoshida in June. In July, Sargassum nipponicum Yendo and Sargassum piluliferum (Turner) C. Agardh dominated. Subsequently, the major species shifted to Sargassum ringgoldianum Harvey and S. yamamotoi in August, Sargassum micracanthum (Kützing) Endlicher, Sargassum macrocarpum C. Agardh and Zostera marina Linnaeus in September, and S. ringgoldianum and S. micracanthum in October. However, the occurrence of S. yamamotoi, S. nipponicum and S. piluliferum in June or July were particularly heterogeneous compared with other areas of Japan. Dendrogram analysis was done based on frequency of appearance. Pooled monthly samples were divided into three groups characterized from the dominant species, degree of domination, weight, length and number of species of drifting seaweeds as well as the degree of diversity or evenness in appearance. This characterization indicated that the diversity and abundance of drifting seaweeds were higher from April to June than in later months.     

Taxonomy and distribution of Sargassum (Phaeophyceae) in the Gulf of Thailand

Ten species of Sargassum (Sargassaceae, Phaeophyceae) were found along the Gulf of Thailand. Morphological characteristics of Sargassum baccularia (Mertens) C.A. Agardh, S. binderi Sonder, S. cinereum J.G. Agardh, S.crassifolium J.G. Agardh, S. longifructum Tseng et Lu, S. oligocystum Montagne, S. polycystum C.A. Agardh, S. siliquosum J.G. Agardh, S. swartzii (Turner) C.A. Agardh and one unidentified species were examined and are described in detail. The most common species were S. polycystum distributed widely in almost all the study sites, S. crassifolium restricted to Prachuap Khirikhan Province, S. longifructum restricted to Chumphon Province, S. siliquosum restricted to Surat Thani Province and one unidentified species restricted to Songkhla Province. Three species (S. cinereum, S. longifructum and S. swartzii) are new records for the algal flora of Thailand. Five species (S. baccularia, S. cinereum, S. longifructum, S. polycystum and the unidentified species) belong to the section Zygocarpicae (J.G. Agardh) Setchell.     

Testing systematic concepts of Sargassum (Fucales, Phaeophyceae) using portions of the rbcLS operon

The taxonomic and phylogenetic concepts within the Sargassum C. Agardh (Sargassaceae) species complex were evaluated through molecular phylogenetic analyses using portions of the chloroplast encoded rbcLS Operon. According to more conservative sequences (rbcL), Turbinaria (Turner) J. Agardh is a close and well‐supported sister lineage to the Sargassum species complex and an appropriate external outgroup for analyses of subgenera and subsections within Sargassum. Both rbcL and more rapidly evolving rbcLS spacer sequences indicated that the East Asiatic genus Myagropsis (Mertens et Turner) Fensholt, along with Sargassum sinicola Setchell et Gardner, represent the closest lineage to Sargassum and form appropriate internal outgroups. The rbcLS spacer region supported three of four subgeneric designations by J. Agardh and sectional levels within the subgenus Sargassum. However, some aspects of Agardh's system were not supported: many of the subsectional ranks or the phyletic concepts; Phyllotrichia was not monophyletic as a subgenus, and its species were also not the most ancestral of Sargassum; and subgenus Sargassum was not the most derived subgenus within the genus. This modern phylogeny suggests a deep evolutionary history for subgenus Sargassum with rapid speciation in closely related subsections and series, and a sister relationship between subgenera Arthrophycus and Bactrophycus.     

In situ growth rates of Sargassum horneri (Fucales,Phaeophyta)1

The relative growth rate of the brown alga Sargassum horneri (Turner) C. Agardh in the sea at Maizuru Bay, Japan, was investigated from summer to winter to examine the impact of declining solar radiation and temperature. The observed change in the growth rate between summer and winter was insignificant, in spite of a major decline in solar radiation and water temperature. The mean relative growth rate was 4.6% per day, equivalent to the daily net photosynthetic production previously reported for this species.     

Further confirmation of the presence of DNA in the pyrenoid core of the siphonous green algal genus Caulerpa (Caulerpales, Ulvophyceae, Chlorophyta)

We re-examined the distribution of chloroplast DNA (ct-DNA) in the pyrenoid core of Caulerpa okamurae Weber van Bosse and C. lentillifera J. Agardh by fluorescence microscopy after staining the squashes and Technovit sections with DNA fluorochromes such as 4′6-diamidino-2-phenylmdole (DAPI), ethidium bromide, Hoechst 33258 and chromomycin A3. All fluorochromes stained specifically the pyrenoid core on the squashes and Technovit sections. In addition, we present new data on the localization of ct-DNA in the pyrenoid core of two other species of the genus Caulerpa: C. cactoides (Turner) Agardh and C. geminata Harvey.     

Phylogenetic relationships of Sargassum (Sargassaceae, Phaeophyceae) with reference to a taxonomic revision of the section Phyllocystae based on ITS-2 nrDNA sequences

Sequences of the end of the 5.8S gene and the internal transcribed spacer 2 (ITS‐2) of nuclear ribosomal DNA have been determined for 19 species of the brown algal genus Sargassum (Sargassaceae), representing three subgenera and eight sections (sections are in parentheses): Phyllotrichia, Bactrophycus (Teretia, Spongocarpus, Halochloa and Repentia) and Sargassum (Acanthocarpicae, Malacocarpicae, Zygocarpicae) to assess the taxonomic position of the section Phyllocystae traditionally included within the Bactrophycus. The sequence of Myagropsis myagroides (Mertens ex Turner) Fensholt (Sargassaceae) was used as an outgroup. Sequences of ITS‐2 were analyzed using neighbor‐joining, parsimony and maximum likelihood methods. The results showed the existence of three clades in Sargassum, corresponding to the three subgenera. The subgenus Phyllotrichia is positioned near the outgroup. Two robust clades were obtained, one corresponding to the subgenus Bactrophycus and the other to the subgenus Sargassum. Sargassum mcclurei Setchell and Sargassum quinhonense Nguyen, the two Phyllocystae investigated, are close to species belonging to the section Zygocarpicae in the subgenus Sargassum. A transfer of the section Phyllocystae to the subgenus Sargassum is therefore proposed on the basis of molecular data (ITS‐2) and morphological data (receptacles and basal leaf).     

DNA SEQUENCE DATA DEMONSTRATE THE POLYPHYLY OF THE GENUS CYSTOSEIRA AND OTHER SARGASSACEAE GENERA (PHAEOPHYCEAE)1

Phylogenetic relationships in the Sargassaceae were explored using three DNA markers, and the monophyly of its genera was challenged. Nineteen out of 24 currently recognized genera were sampled, representing 63 species. The variable mt23S‐tRNA Val intergenic spacer could only be aligned within genera and could not be used to infer intergeneric relationships. The partial mt23S was also useful to delineate genera and was alignable at the family level but provided few informative characters. Analysis of mt23S DNA sequences together with chloroplast‐encoded psbA sequences resulted in a better resolved phylogeny. Hormophysa was the first genus to branch off within the Sargassaceae, followed by Myriodesma; then the three genera Caulocystis, Carpoglossum, and Scaberia in unresolved order; and then Acrocarpia. The other taxa studied here were divided over three major clades, but there was no branch support for the monophyly of two of these. The genera Bifurcaria, Cystoseira, Halidrys, and Sargassum appeared polyphyletic. The following taxonomic changes are proposed: a new genus Brassicophycus for Bifurcaria brassicaeformis (Kützing) E. S. Barton; reinstatement of the genus Sargassopsis for Sargassum decurrens (R. Brown ex Turner) C. Agardh; reinstatement of the genus Sirophysalis for Indo‐Pacific Cystoseira trinodis (Forsskål) C. Agardh; reinstatement of the genus Polycladia for the western Indian Ocean species Cystoseira indica (Thivy et Doshi) Mairh, Cystoseira myrica (S. G. Gmelin) C. Agardh, and Acystis heinii Schiffner; and reinstatement of the genus Stephanocystis for the North Pacific Cystoseira species and Halidrys dioica N. L. Gardner. The European Cystoseira species should be split into three genera, but no name changes are proposed yet, because diagnostic characters were found only for the clade including the type species. Some evolutionary trends could be discerned from the mt23S + psbA phylogeny.     

Long-Term Biomonitoring of Heavy Metal Pollution of Ussuri Bay,Sea of Japan

Russian Journal of Marine Biology - The concentrations of Fe, Mn, Cu, Zn, Pb, Cd, and Ni were studied in the brown algae Sargassum miyabei Yendo and S. pallidum (Turner) C. Agardh collected in...     

EVALUATION OF RECOVERABLE FUNCTIONAL LIPID COMPONENTS OF SEVERAL BROWN SEAWEEDS (PHAEOPHYTA) FROM JAPAN WITH SPECIAL REFERENCE TO FUCOXANTHIN AND FUCOSTEROL CONTENTS1

Fucoxanthin (Fx) and fucosterol (Fs) are characteristic lipid components of brown seaweeds that afford several health benefits to humans. This article describes the quantitative evaluation of lipids of 15 species of brown seaweeds with specific reference to Fx, Fs, and functional long‐chain omega‐6/omega‐3 polyunsaturated fatty acids (PUFAs). In addition, fatty‐acid composition of selected species was also accomplished in the study. Major omega‐3 PUFAs in the brown seaweeds analyzed were α‐linolenic acid (18:3n‐3), octadecatetraenoic acid (18:4n‐3), arachidonic acid (20:4n‐6), and eicosapentaenoic acid (20:5n‐3). Both Fx (mg · g^?1 dry weight [dwt]) and Fs (mg · g^?1 dwt) were determined to be relatively abundant in Sargassum horneri (Turner) C. Agardh (Fx, 3.7 ± 1.6; Fs, 13.4 ± 4.4) and Cystoseira hakodatensis (Yendo) Fensholt (Fx, 2.4 ± 0.9; Fs, 8.9 ± 2.0), as compared with other brown seaweed species. Studies related to seasonal variation in Fx, Fs, and total lipids of six brown algae [S. horneri, C. hakodatensis, Sargassum fusiforme (Harv.) Setch., Sargassum thunbergii (Mertens ex Roth) Kuntze, Analipus japonicus (Harv.) M. J. Wynne, and Melanosiphon intestinalis (D. A. Saunders) M. J. Wynne] indicated that these functional lipid components reached maximum during the period between January and March. The functional lipid components present in these seaweeds have the potential for application as nutraceuticals and novel functional ingredients after their recovery.     

THE LIFE CYCLE OE SARGASSUM HORNERI (PHAEOPHYTA) IN LABORATORY CULTURE1

The life cycle of the large dioecious alga Sargassum horneri (Turner) C. Agardh was completed in unialgal culture by controlling photoperiod in relation to the phase of growth. Embryos isolated from a naturally grown female thallus gave rise to early germlings that rapidly formed blades under both short-day (9 h L) and long-day (15 h L) conditions at 20° C Shoot elongation, which followed early blade formation, occurred under the short-day conditions hut not under the long-day conditions. Functional female and male receptacles developed when thalli 8–14 cm long grown under the short-day conditions were transferred to the long-day conditions; gamete fusion occurred when male and female thalli were grown together. Fertilized oospores gave rise to normal thalli in a manner similar to that for in situ plants. Thus, the life cycle of S. horneri was completed in laboratory culture.     

MORPHOLOGICAL AND GENETIC VARIATION IN THE POPULATIONS OF SARGASSUM HEMIPHYLLUM (PHAEOPHYCEAE) IN THE NORTHWESTERN PACIFIC1

Difficulty in species identification of Sargassum (Sargassaceae, Fucales) is partly attributed to the high polymorphism among its individuals and populations. This study aimed at assessing morphological and genetic variations in two varieties, var. hemiphyllum J. Agardh and var. chinense J. Agardh, of Sargassum hemiphyllum (Turner) C. Agardh, a widely distributed species in the northwestern Pacific. We investigated 26 measurable, five numerical, and 33 categorical morphological parameters associated with different branching levels of specimens from each of six localities within its distribution range using cluster analysis (CA) and principal coordinate analysis (PCoA). Leaf size of the primary and secondary branching levels and the vesicle size of the secondary branches of the specimens examined were determined to be the most important morphological parameters that were significantly different among populations. Change in leaf and vesicle length of individuals among the six populations followed a latitudinal gradient, with smaller leaves and vesicles associated with northern populations and larger ones in the southern populations. The possible influence of the gradual change in sea surface temperatures (SSTs) along this gradient in the northwestern Pacific on leaf and vesicle morphologies of this species was suggested. PCR‐RFLP analysis of the RUBISCO spacer in the chloroplast genome revealed two distinct and highly homogenous clades, a China clade and a Japan‐Korea clade, which corresponded to var. chinense and var. hemiphyllum, respectively. The formation of refugia along the “Paleo‐coast” in the East China Sea during glacial periods is suggested to have led to the vicariance of ancestral populations of S. hemiphyllum and thus to have promoted genetic differentiation. The massive freshwater outflow of the Yellow and Yangtze rivers may continue to act as a barrier, prolonging the allopatric distribution of the two varieties.     

Phylogenetic relationships within the genus Sargassum (Fucales, Phaeophyceae), inferred from ITS-2 nrDNA, with an emphasis on the taxonomic subdivision of the genus

Sequences from the ribosomal DNA internal transcribed spacer‐2 (ITS‐2) were compared among species of Sargassaceae including the genera Sargassum and Hizikia. Species of different subgenera and sections of Sargassum were used to assess the taxonomic relationships within the genus, especially the subdivisions of the subgenus Bactrophycus. Sequences were aligned in accordance with their common secondary structure. Phylogenetic trees were constructed using neighbor‐joining, maximum likelihood and maximum parsimony methods with three species of Turbinaria as outgroups. The resulting phylogenetic trees showed that the genus Sargassum is divided into three clades corresponding to the subgenera Phyllotrichia, Sargassum and Bactrophycus. This last subgenus is further divided into four distinct groups: a Spongocarpus clade, a Teretia clade, a Hizikia clade, and a Halochloa/ Repentia clade. The position of the section Phyllo‐cystae, excluded from the subgenus Bactrophycus and included within the subgenus Sargassum is once again confirmed by the present study. Current results strongly support the assignation of Hizikia fusiformis to the genus Sargassum. Based on morphological differences and a distinct position in the molecular trees, Hizikia should be recognized as a section in the subgenus Bactrophycus so that Hizikia (Okamura) Yoshida, stat. nov. is proposed. A remarkably low divergence of ITS‐2 sequences was observed for the species in the sections Repentia and Halochloa, suggesting very recent radiation of these species. The subgenus Sargassum is divided into three clades corresponding to the three known sections: Acanthocarpicae, Malacocarpicae and Zygocarpicae, previously recognized by the morphology of receptacles. The position of Sargassum duplicatum, S. carpophyllum, S.yendoi, S. piluliferum and S. patens within the subgenus Sargassum is discussed.     

Use of RAPD in differentiation of selected species of <Emphasis Type="Italic">Sargassum</Emphasis> (Sargassaceae,Phaeophyta)

Sargassum C. Agardh is one of the most common but little understood genera of Phaeophyta in Malaysia. The difficulty in species delineation is due to morphological plasticity. A combination of morphology and random amplified polymorphic DNA (RAPD) studies of selected Sargassum species was carried out to have a better understanding of the taxonomy. Primer OPA13 was found to be good for discriminating between Sargassum species. Sargassum binderi was shown to be different from S. oligocystum (S_D>0.5 = 14.11%), indicating the importance of the vesicle and receptacle in species differentiation. S. baccularia was clearly separated out from S. polycystum and S. stolonifolium using primer OPA13. RAPD analysis showed that the presence of the stolon is an important character for separating S. baccularia (no stolon) from S. polycystum (stolon) and S. stolonifolium (stolon). Presented at the 6th Meeting of the Asian Pacific Society of Applied Phycology, Manila, Philippines.     

ITS1 sequences of type specimens of Gigartina and Sarcothalia and their significance for the classification of South African Gigartinaceae (Gigartinales,Rhodophyta)

Uncertainties about the identity of type specimens of red algae have frequently led to taxonomic and nomenclatural confusion. A procedure for extracting PCR-amplifiable DNA from formalin-fixed material and herbarium specimens was used to investigate the taxonomic status of several South African Gigartinaceae. We compared nucleotide sequences in the internal transcribed spacer 1 (ITS1) region in type specimens and other historically important collections presently referred to Gigartina bracteata, G. radula and G. stiriata. The following opinions are supported: (1) Fucus bracteatus S.G. Gmelin, Chondrodictyon capense Kützing and Iridaea clathrata Decaisne represent a single species of South African Gigartina (G. bracteata (S.G. Gmelin) Setchell & Gardner) in which the disintegration of tetrasporangial sori results in a reticulate thallus. (2) Mastocarpus polycarpus Kützing, M. incrassatus Kützing and Iridaea lapathifolia Kützing represent a single species of South African Gigartina (G. polycarpa (Kützing) Setchell & Gardner) that has often, but erroneously, been called G. radula (Esper) J. Agardh. (3) Mastocarpus verrucosus Kützing is a later heterotypic synonym of Iridaea papillosa Bory (Sarcothalia papillosa (Bory) Leister) and was based on material that probably came from southern South America rather than from South Africa, the provenance given by Kützing. (4) Fucus stiriatus Turner and Sphaerococcus burmannii C. Agardh represent a single species of South African Sarcothalia (S. stiriata (Turner) Leister).     

SEASONAL CHANGES IN SIZE STRUCTURE OF SARGASSUM AND TURBINARIA POPULATIONS (PHAEOPHYCEAE) ON TROPICAL REEF FLATS IN THE SOUTHERN RED SEA1

Seasonal variation in density, thallus length and biomass, population size structure, and allometric length‐biomass relationships was investigated in populations of Sargassum ilicifolium (Turner) C. Agardh, Sargassum subrepandum (Forssk.) C. Agardh, and Turbinaria triquetra (J. Agardh) Kütz. (Phaeophyceae) on shallow reef flats in the southern Red Sea. Thallus length and biomass varied strongly with season, with the highest values occurring in the cooler months. Thallus densities showed no significant temporal variation. Log‐total biomass versus log‐density relationships were positive throughout the growth season without any decrease in the slope of the relationship. In two populations, biomass‐density combinations approached the interspecific biomass‐density line, but the massive annual shedding of modules occurred before self‐thinning would set in. Allometric length‐biomass relationships varied with season in all populations and were associated with seasonal module initiation, growth, and shedding. Evidence of a strong asymmetric competition was found in two high‐density populations. These populations showed a predominance of small thalli during peak development, asymmetrical Lorenz curves, increasing Gini coefficients, and increasing thallus length relative to biomass during the main growth phase. In two other less crowded populations, small thalli were absent during peak development, Lorenz curves were symmetrical, and Gini coefficients decreased during the main growth phase. In these populations, size equalization appears to be due to responses at the modular level rather than size‐dependent mortality. We conclude that changes in size structure in this highly seasonal environment are determined by module dynamics, modified by asymmetric competition in some populations, with a minor role of recruitment and no regulatory effect of self‐thinning.     

PHYLOGENETIC RELATIONSHIPS WITHIN THE GENUS HYPNEA (GIGARTINALES,RHODOPHYTA), WITH A DESCRIPTION OF H. CAESPITOSA SP. NOV.1

Species discrimination within the gigartinalean red algal genus Hypnea has been controversial. To help resolve the controversy and explore phylogeny within the genus, we determined rbcL sequences from 30 specimens of 23 species within the genus, cox1 from 22 specimens of 10 species, and psaA from 16 species. We describe H. caespitosa as a new species characterized by a relatively slender main axis; a pulvinate growth habit with entangled, anastomosing, and subulate uppermost branches; and unilaterally borne tetrasporangial sori. The new species occurs in the warm waters of Malaysia, the Philippines, and Singapore. The phylogenetic trees of rbcL, psaA, and cox1 sequences showed a distant relationship of H. caespitosa to H. pannosa J. Agardh from Baja California and the marked differentiation from other similar species. The rbcL + psaA tree supported monophyly of the genus with high bootstrap values and posterior probabilities. The analysis revealed three clades within the genus, corresponding to three sections, namely, Virgatae, Spinuligerae, and Pulvinatae first recognized by J. G. Agardh. Exceptions were H. japonica T. Tanaka in Pulvinatae and H. spinella (C. Agardh) Kütz. in Spinuligerae.