Systematic studies of the antarctic species of the phyllophoraceae (Gigartinales,Rhodophyta) based on rbcL sequence analysis

The taxonomic placement of four antarctic species of the marine red algal family Phyllophoraceae (Gigartinales) is assessed within a preliminary molecular phylogeny of the family based on direct sequence analysis of the chloroplast gene rbcL. Parsimony analysis of rbcL sequences indicates that Gymnogongrus antarcticus and Gymnogongrus turquetii cluster in a clade consisting predominantly of southern hemisphere species currently placed in Gymnogongrus and Ahnfeltiopsis, whereas Phyllophora ahnfeltioides and Phyllophora antarctica cluster in a separate clade that is widely divergent from the northern hemisphere Phyllophora clade. Results from molecular and morphological data challenge the current taxonomic concept that type of life history is a phylogenetically valid criterion for recognition of genera in the Phyllophoraceae.     

Molecular Systematics and Phylogenetics of Gracilariacean Species from the Mediterranean Sea

Mediterranean reports of Gracilariaceae species, in particular those assigned to the G. verrucosa complex, were re-examined with the use of molecular tools, in order to verify their systematic position and better understand their distribution. Within this complex, we recognized four distinct taxa: Gracilariopsis longissima, Gracilaria gracilis, Gracilaria longa and a possible new species. The rbcL gene sequences, together with those of other terete Mediterranean entities, were included in a broad molecular phylogeny of the family. The reproductive characters of the studied taxa do not fit completely with published hypothesis on the generic and intrageneric relationships, suggesting that the anatomy of some subgroups should be better characterized.     

Taxonomic Considerations of a Foliose Grateloupia Species from the Straits of Messina

Grateloupia turuturu Yamada is the currently accepted name for the invasive red alga that is present on coasts of the North Atlantic. Previously considered as G. doryphora (Montagne) M.A. Howe, populations of this invasive species were examined and their taxonomic position revised using molecular and morphological techniques. It was also thought that similar invasive populations in the Mediterranean should be identified as G. turuturu. This investigation used rbcL based molecular analyses to clarify the taxonomic position of Grateloupia “doryphora’ from the Straits of Messina. Our results indicate that this population is neither G. doryphora nor G. turuturu. It was placed separately in all analyses and grouped consistently with other Grateloupia species from the Pacific. On the basis of molecular data from this and previous investigations, it is evident that the status of the foliose Atlantic and Mediterranean entities is still unclear and a re-evaluation of the old names connected to them should be undertaken.     

A simple and rapid technique to PCR amplify plastid genes from spores of <Emphasis Type="Italic">Porphyra</Emphasis> C. Agardh (Bangiales,Rhodophyta)

A user-friendly and fast protocol using PCR was developed to amplify the plastid rbcL gene of Porphyra, the most economically valuable genus of Rhodophyta. The technique involved the use of monospores, providing the advantage of a small and genetically homogeneous set of cells and thus reducing the risk of biological or chemical contamination. An additional factor in the choice of the rbcL gene was the huge amount of sequence data deposited in molecular biology databases and their utility as molecular markers in taxonomic studies.     

Evidence for a composite phylogenetic origin of the plastid genome of the brown alga Pylaiella littoralis (L.) Kjellm

The nucleotide sequence and the 5 flanking region of the rbcL gene coding for the large subunit of ribulose bisphosphate-1,5-carboxylase/oxygenase of Pylaiella littoralis, a brown alga, has been determined and the deduced amino-acid sequence has been compared to those of various photosynthetic and chemoautotrophic Eubacteria, of a red alga and of green plastids (Euglena gracilis, green algae and higher plants). Unlike the rbcL genes of green plastids which are more closely related to those of cyanobacteria the P. littoralis rbcL gene is more closely related to that of a -purple bacterium, as was found for the rbcS gene of another chromophytic alga [Boczar et al., Proc Natl Acad Sci USA 86: 4996–4999, 1989]. Matrix data of homology between the rbcL gene of P. littoralis and the same gene of other organisms are presented. Based on our previous report, the gene coding for the 16S rRNA from P. littoralis is closely related to that of E. gracilis (Markowicz et al., Curr Genet 14: 599–608, 1988). We suggest that the large plastid DNA molecule of P. littoralis is a phylogenetically composite genome which probably resulted from mixed endosymbiosis events, or from a horizontal transfer of DNA.     

Worldwide distribution of commercial resources of seaweeds including Gelidium

The commercial exploitation of seaweeds for use as food and for the production of agar, alginate and carrageenan is outlined. The quantities of seaweed harvested for each purpose are tabulated and discussed. Seaweeds for food are derived chiefly from China, Japan and Korea, with almost 94% obtained by cultivation. Alginophytes are collected in 15 countries but six of these account for more than 80% of the total harvest; all are from natural stocks except for a large quantity of Laminaria cultivated in China. Natural carrageenophytes, from 12 countries, now account for only 20% of the total harvest; the remainder is cultivated Eucheuma species, 99% of which is produced in only two countries, the Philippines and Indonesia. Of the four categories of commercial resources of seaweeds considered, agarophytes are spread more evenly over a greater number of countries; they come from 20 countries and only five of these are minor contributors to the total. Gelidium species are particularly important because of the high quality agar they yield; their distribution and location are discussed.     

Gelidium pristoides in South Africa

Gelidium pristoides has been harvested commercially from the eastern Cape, South Africa, since 1951, with 40–80 t y^–1 (dry wt) collected in recent years. This species has been intensively studied since 1983, and we briefly review knowledge of its biology in relation to harvesting. We describe a new study of intertidal epiphytic animals, showing that none is specific to G. pristoides, and that only 2.8% of these animals (numbers) inhabit this agarophyte, while the rest are found in other intertidal algal communities: harvesting is considered to have negligible effects on epifauna. Over the past 3 y, we have monitored, at two sites, the effects of the harvesting of G. pristoides on other benthic algae and animals. In only two of the seven main components analysed, did we find any difference between harvested and control plots. At one site only, the number of limpets and percentage cover of Gelidium was higher in harvested plots. These results show that harvesting has no significant biological effect. Regulations governing seaweed exploitation in South Africa were amended in 1988, to encourage local processing of products, and these changes are discussed in relation to the local Gelidium industry. Despite experimental results predicting a higher yield per unit effort if harvesting is limited to summer, harvesting continues throughout the year for practical reasons.     

Phylogenetic relationships of the enigmatic fern families Hymenophyllopsidaceae and Lophosoriaceae: Evidence fromrbcL nucleotide sequences

Nucleotide sequences fromrbcL were used to infer relationships of Lophosoriaceae and Hymenophyllopsidaceae. The phylogenetic positions of these two monotypic fern families have been debated, and neither group had been included in recent molecular systematic studies of ferns. Maximum parsimony analysis of our data supported a sister relationship betweenLophosoria andDicksonia, and also betweenHymenophyllopsis andCyathea. Thus, both newly-examined families appear to be part of a previously characterized and well-supported clade of tree ferns. The inferred relationships ofLophosoria are consistent with most (but not all) recent treatments. However,Hymenophyllopsis includes only small delicate plants superficially similar to filmy ferns (Hymenophyllaceae), very different from the large arborescent taxa. Nevertheless, some synapomorphic characteristics are shared with the tree fern clade. Further studies on gametophytes ofHymenophyllopsis are needed to test these hypotheses of relationship.     

Seaweeds of potential economic importance in Kenya: field survey and future prospects

Kenya is a net importer of agar and alginate based on recent government statistics, although it may have the potential to be self-sufficient or even an exporter of these phycocolloids. There is little information on carrageenan importation into Kenya since government statistics incorporate it as agar. Seaweeds are relatively unimportant in the Kenyan diet since they are consumed rarely by coastal people. A survey of 15 sites along of the Kenyan coast evaluated the potential for harvesting seaweeds and for establishing seaweed farms. Gracilaria appears to comprise the bulk of the low grade agar import, even though local species of this genus are widely distributed along the Kenyan coast. Major populations of Gelidium may be a potential source of high quality bacteriological grade agar. Eucheuma may be farmed locally to support the increasing local and regional demands for carrageenan. Recommendations for a national program of management and production for Kenya will be discussed.     

Ethnobotany of seaweeds: clues to uses of seaweeds

Extensive uses of fresh and dried seaweeds by coastal populations over the world can furnish clues to potential food and other uses of seaweeds, just as the first extraction of a seaweed gelatin now used for bacteriological purposes was discovered by a housewife in search of a pudding. Ethnic uses as food depend heavily on closely related species suitable for making cool, gelatinous dishes or concoctions, or on species suitable for adding to soups or stews. Rarely, single species like cochajugo (Durvillea antarctica) in Chile and rimu (Durvillea antarctica) in New Zealand point to different kinds of food preparation. Oriental cuisine incorporates many different species of seaweeds in a wide variety of ways, whereas commercial western uses depend upon extracts of wall materials to suspend, emulsify or stabilize a broad variety of foods and products.     

Recent developments in the systematics of the Gigartinaceae (Gigartinales, Rhodophyta) based on rbcL sequence analysis and morphological evidence

The phylogenetic systematics of the Gigartinaceae is discussed for seven genera and three undescribed generic lineages and 65 taxa representing 62 species based on an analysis of rbcL sequences and morphological evidence. An examination of rbcL trees resulting from analyses of these taxa identifies seven lineages: (i) ‘Gigartina’ alveata; (ii) Rhodoglossum/Gigartina; (iii) Chondracanthus; (iv) Ostiophyllum; (v) Sarcothalia; (vi) ‘Gigartina’ skottsbergii; and (vii) a large clade containing Iridaea/‘Sarcothalia’, Mazzaella and Chondrus. These lineages and Chondrus are strongly supported; however, two groups, Iridaea/‘Sarcothalia’ and Mazzaella, receive no bootstrap support. The morphology of the female reproductive system is investigated with the aid of computer-generated, color-coded tracings of photographs of cystocarps seen in cross section at different developmental stages. Seven basic cystocarp types were found which corresponded to species groups seen in rbcL trees. These were: (i) a ‘Gigartina’ alveata group in which the carposporangia-bearing filaments develop apomictically from gametophytic cells; (ii) a Rhodoglossum/Gigartina group in which gonimoblast filaments penetrate the surrounding envelope fusing progessively with envelope cells; (iii) a Chondracanthus group in which gonimoblast filaments penetrate the envelope but fuse with envelope cells only at late developmental stages; (iv) a Sarcothalia group in which the gonimoblast filaments displace an envelope composed mainly of secondary gametophytic filaments and link to envelope cells by terminal tubular gonimoblast cells; (v) an Iridaea group similar to the Sarcothalia group, but with an envelope composed of a mixture of medullary cells and secondary gametophytic filaments; (vi) a Mazzaella group that lacks a true envelope and in which gonimoblast filaments connect to modified gametophytic cells by means of terminal tubular cells; (vii) a Chondrus group in which gonimoblast filaments penetrate the medulla and link to modified medullary cells by means of conjunctor cells forming secondary pit connections. The further separation of these groups into genera is based largely on tetrasporangial characters.     

Phylogenetic analysis ofEremosyne pectinata (Saxifragaceae s. l.) based onrbcL sequence data

Earlier analyses ofrbcL sequences clarified phylogenetic relationships of 16 of 17 subfamilies ofSaxifragaceae s. l. This study investigates the affinities of the monotypicEremosynoideae, the only subfamily ofSaxifragaceae s. l. not examined previously forrbcL sequence variation. Our analyses suggest thatEremosyne is only distantly related to core members ofSaxifragaceae s. l. (i.e.Saxifragoideae = Saxifragaceae s. str.); it is allied instead with members ofAsteridae. A particularly close relationship is indicated betweenEremosyne andEscallonia, another traditional member ofSaxifragaceae s. l. that also appears as part ofAsteridae. Inclusion ofEremosyne withinAsteridae is in agreement with embryology, as well as with 18S rDNA sequence data.     

Systematic position of Asteropyrum (Ranunculaceae) inferred from chloroplast and nuclear sequences

Sequences of chloroplast gene rbcL and partial nuclear 26S rDNA were used to evaluate phylogenetic relationships of Asteropyrum. Four primary clades were recognized in Ranunculaceae, corresponding to subfamilies Hydrastidoideae, Coptidoideae, Thalictroideae, and Ranunculoideae. Our results place Asteropyrum in Ranunculoideae, sister to the tribe Actaeeae, which includes Beesia, Cimicifuga, and Eranthis. This is supported by chromosome characters, including chromosome size and basic number, and the stainability of prophase chromosomes and interphase nuclei. Our results do not support previous placements of Asteropyrum in either Coptidoideae or Thalictroideae. Considering its uniqueness in a few characters (e.g. simple peltate leaves, accumulating benzylisoquinoline alkaloids, vessel elements with only typical scalariform perforation plates), we recognize Asteropyrum as a monotypic tribe of Ranunculoideae, Asteropyreae W. T. Wang et C. Y. Chang.     

Patterns of gametophyte dominance ofIridaea splendens(Rhodophyta) in Vancouver Harbour,Vancouver, British Columbia,Canada

This paper presents results of field and laboratory work on one component of population structure ofIridaea splendens (Setchellet Gardner) Papenfuss (Gigartinaceae, Rhodophyta), the seasonal change in abundance of the gametophyte life-history stage. Investigations of this genus by others (using a variety of sampling and identification techniques) have shown a seasonal gametophyte dominance, sporophyte dominance, or some combination of these two. Gametophyte stages ofIridaea splendens produce kappa carrageenan and tetrasporophyte stages lambda carrageenan; a chemical colorimetric test using resorcinol can be used to identify these stages regardless of the presence of reproductive structures. In this paper we report on the proportion of gametophytes determined both by this chemical test and by the presence of reproductive structures, and on one possible determinant of the seasonal change in dominance. Analysis of field samples using the resorcinol test showed that from June–August the gametophyte stage predominates and from December–February the tetrasporaphyte stage is most common, both in reproductive and non-reproductive thalli (the latter tested as three size classes). Examination of reproductive structures gave similar results. One factor suggested in the literatures as a possible determinant of the haploid/diploid ratio is apomeiosis (in the formation of the tetraspores); our results show this to be unimportant in the Vancouver Harbor populations.Presented at the XIIIth International Seaweed Symposium, University of British Columbia, Vancouver, Canada, August 1989.     

Chemical screening of Brazilian marine algae producing carrageenans

The yield, IR spectra, 3,6-anhydro-d-galactose and sulfate content are given for several species of putative carrageenophytes from different sites on the Brazilian coast. Kappa carrageenan was found in Hypnea musciformis and Gigartina spp. ; all the other species tested yield an iota type. The lowest carrageenan yield was found in Cryptonemia crenulata (5%), and the highest in Gigartina sp. (72%); 3,6-anhydrogalactose content was highest in Hypnea musciformis and Gigartina spp., whereas sulfate was lowest in H. musciformis. Slight variations in the yield of carrageenans were found during the period of observation (March 1988 to May 1989), but no clear pattern could be recognized. No significant variation was observed in plants from different collection sites.     

Temporal and spatial variation in agar from a population of Pterocladia capillacea (Gelidiales,Rhodophyta) from Brazil

Pterocladia capillacea forms dense intertidal belts in southeastern Brazil, on moderately exposed rocky coasts. The studied population extends along a gradient of water exposure, where slightly different morphotypes can be recognized. Specimens were collected monthly from 3 points along the exposure gradient of its distribution (lower, medium and higher exposure), and analyzed for agar, sulfate and 3,6 anhydrogalactose content. Agar varied from 5–32% of dried seaweed with lower yields in the winter, and higher yields in late spring/early summer. Specimens from the surf side of the distribution had a consistently higher agar content throughout the year. Sulfate varied from 1–5%, and 3,6 AG from 27–48% of dried agar, without a clear variation among the sites.     

The effect of phosphate concentration on growth and agar content of Gelidium robustum (Gelidiaceae,Rhodophyta) in culture

Phosphate concentration of the growth medium was found to affect the growth rate and agar yield of a clone of Gelidium robustum grown in the laboratory. To study differences in growth we used phosphate concentrations from 0 to 200 µM. To determine the effect of phosphate on agar yield and its properties we used concentrations from 0 to 20 µM. Growth rates generally increased with increasing phosphate concentration, with the highest growth rate (21% d^–1) obtained at 150 µM. Agar yield as percentage of fresh weight was highest (10%) in the algae grown with low phosphate concentrations, but agar yield as percentage of dry weight was highest(43%) at 20 µM of phosphate. Gel strength increased with phosphate concentration with a maximum of 160 g m^–2 for 0.75% gels for the cultures at 20 µM. Melting and gelling temperatures of the gels were also affected by phosphate concentration of the growth medium. Starch yield was highest in algae grown in low phosphate concentrations.     

Interspecific Relationships among Charrs Based on Phylogenetic Analysis of Nuclear Growth Hormone Intron Sequences

DNA samples were obtained from six charr species: Salvelinus alpinus, S. malma, S. confluentus, S. leucomaenis, S. namaycush and S. fontinalis and two outgroups: Salmo salar and S. trutta. The C and D introns from the duplicate growth hormone loci (GH1 and GH2) were amplified using the polymerase chain reaction and sequenced from each of the species. Phylogenetic analyses with and without gaps were performed using the beta version of PAUP. The GH introns are evolving relatively slowly, so the data were combined with sequences from the ITS1 and ITS2 for a phylogenetic analysis of the charr species. The GH intron data and the combined data supported two groups of sister species among charrs: S. alpinus and S. malma, and S. confluentus and S. leucomaenis.