UNUSUAL PHENOMENA IN THE LIFE HISTORIES OF FLORIDEAE IN CULTURE1

Several Florideae grown in natural seawater media under defined laboratory conditions have interesting and unusual life histories. Antithamnion occidentale males of one generation produced tetraspores that gave rise to nonsporangiate males. The functional females of A. pygmaeum developed spermatangia and tetrasporangia; the tetraspores formed new females. Antithamnion defectum tetrasporophytes of one generation bore spermatangia in addition to tetrasporangia; the tetraspores gave rise to typical gametophytes. Tetraspores from successive generations of Callitham-nion sp. developed into tetrasporophytes and males but no females were produced. Functional female gametophytes of Platythamnion sp. bore abortive tetrasporangia. Field-collected plants of two species of Fauchea produced tetraspores that yielded additional sporangiate plants: those of F. pygmaea being bispo-rangiate and tetrasporangiate, and those of F. lacini-ata being strictly tetrasporangiate. Male plants of Pleonosporium vancouverianum from a running seawater table bore spermatangia and polysporangia when collected. The same plants in unialgal culture produced only spermatangia.     

Correlations between primary and secondary metabolites in Ceramiales (Rhodophyta)

A correlation between the presence of terpenes and floridosides and between the presence of bromophenols and digeneaside was observed in species of red algae belonging to the order Ceramiales. This observation suggests a division in Ceramiales that can be easily detected through the presence of the secondary metabolites, terpenes and bromophenols.     

Host specificity and growth of kelp gametophytes symbiotic with filamentous red algae (Ceramiales,Rhodophyta)

Kelp gametophytes were previously observed in nature living endophytically in red algal cell walls. Here we examine the interactions of two kelp species and six red algae in culture. Gametophytes of Nereocystis luetkeana (Mertens) Postels et Ruprecht became endophytic in the cell walls of Griffithsia pacifica Kylin and Antithamnion defectum Kylin, and grew epiphytically in high abundance on G. japonica Okamura and Aglaothamnion oosumiense Itono. Alaria esculenta (Linnaeus) Greville from the Atlantic coast of Nova Scotia became endophytic in Aglaothamnion oosumiense, Antithamnion defectum, Callithamnion sp., G. japonica, G. pacifica, and Pleonosporium abysicola Gardner, all from the Pacific Ocean. Some cultures were treated with phloroglucinol before infection to thicken the cell walls. The endophytic gametophytes were smaller and grew more slowly than gametophytes epiphytic on the same host. N. luetkeana failed to become endophytic in some of the potential hosts, and this may reflect host specificity, or culture artifacts. This work improves our understanding of the process of infection of red algae by kelp gametophytes, and broadens our knowledge of host specificity in endophytic symbioses.Communicated by K. Lüning     

PHYLOGENETIC UTILITY OF rbc S SEQUENCES: AN EXAMPLE FROM ANTITHAMNION AND RELATED GENERA (CERAMIACEAE, RHODOPHYTA)

We report the potential phylogenetic utility of the small RUBISCO subunit ( rbc  S) sequences from a sampling of Antithamnion and related genera in ceramiacean algae. The size of rbc  S was 417 bp for all taxa examined. Analyses of the DNA sequence data indicated that pairwise divergences of rbc  S sequences were 3.3%–9.8% among species of Antithamnion , and ranged from 13.6% to 18.0% between Antithamnion and related genera. Phylogenetic analyses fully resolved relationships at the intrageneric level with statistical significance supported by high bootstrap values. Two subgenera of Antithamnion , Pteroton and Antithamnion , were clearly distinguished in the molecular tree. In the clade of subgenus Pteroton , A. aglandum was allied with A. callocladum and separated from A. nipponicum. In the pairwise distance comparison of sequence variation, Ceramium showed the greatest genetic distances among genera examined in the study. All phylogenetic trees generated by the maximum parsimony, neighbor joining, and maximum likelihood were completely congruent in topology with high confidence.     

Gamete recognition during fertilization in a red alga,Antithamnion nipponicum

Summary Fertilization in the marine red algaAntithamnion nipponicum is a highly specific process involving non-motile male gametes, spermatia, and female receptive structures, carpogonia. FITC-lectin and Calcofluor white ST labelling show that the outer cell walls of spermatia differ from vegetative cells in carbohydrate composition. Specific binding of the lectins to spermatial walls was confirmed by lectin-gold labelling on thin sections. Gametic recognition inAntithamnion nipponicum is based on the interaction of a surface carbohydrate on the spermatia with a surface carbohydrate receptor on the trichogynes. Spermatial binding to trichogynes is inhibited by pre-incubation with concanavalin A and trichogyne receptors are blocked by the complementary carbohydrate -D-methyl mannose. The inhibitory effects of concanavalin A to spermatial binding of trichogynes is reversed by preincubation with -D-methyl mannose. The combination of long spermatial appendages and a carbohydrate-carbohydrate receptor-based gamete recognition mechanism make fertilization in this species an efficient process.     

Occurrence of conjugated polyenoic fatty acids in seaweeds from the Indian Ocean

Three species of red marine macro algae (Rhodophyta) from the Indian Ocean were analysed for the occurrence of conjugated polyenes. The composition of different lipid classes in these seaweeds along with their fatty acid composition has also been reported. Analysis of lipid classes of these seaweeds revealed that both Acanthophora spicifera (Ceramiales, Rhodophyta) and two species of Gracilaria, viz. G. edulis and G. folifera (Gracilariales, Rhodophyta) were rich in glycolipids followed by neutral- and phospholipids. The fatty acid composition of these seaweeds revealed C16:0 as the predominant fatty acid in all three species. However, A. spicifera had significantly higher amounts of eicosapentaenoic acid (EPA) and arachidonic acid (AA) as compared to negligible amount of these fatty acids in both species of Gracilaria. The red seaweed Acanthophora spicifera contained conjugated eicosapentaenoic acid (CEPA) and conjugated arachidonic acid (CAA) in all lipid classes except glycolipids.     

Assessing the use of the mitochondrial cox1 marker for use in DNA barcoding of red algae (Rhodophyta)

The red algae, a remarkably diverse group of organisms, are difficult to identify using morphology alone. Following the proposal to use the mitochondrial cytochrome c oxidase subunit I (cox1) for DNA barcoding animals, we assessed the use of this gene in the identification of red algae using 48 samples plus 31 sequences obtained from GenBank. The data set spanned six orders of red algae: the Bangiales, Ceramiales, Corallinales, Gigartinales, Gracilariales and Rhodymeniales. The results indicated that species could be discriminated. Intraspecific variation was between 0 and 4 bp over 539 bp analyzed except in Mastocarpus stellatus (0-14 bp) and Gracilaria gracilis (0-11 bp). Cryptic diversity was found in Bangia fuscopurpurea, Corallina officinalis, G. gracilis, M. stellatus, Porphyra leucosticta and P. umbilicalis. Interspecific variation across all taxa was between 28 and 148 bp, except for G. gracilis and M. stellatus. A comparison of cox1 with the plastid Rubisco spacer for Porphyra species revealed that it was a more sensitive marker in revealing incipient speciation and cryptic diversity. The cox1 gene has the potential to be used for DNA barcoding of red algae, although a good taxonomic foundation coupled with extensive sampling of taxa is essential for the development of an effective identification system.     

Concise review of the genus Vertebrata S.F. Gray (Rhodophyta: Ceramiales)

Journal of Applied Phycology - The red algal genus Vertebrata (Ceramiales, Rhodophyta) comprises 30 species of rather small filamentous algae, differing in morphology, distribution, and ecological...     

Rhodolith beds at the easternmost extreme of South America: Community structure of an endangered environment

Rhodolith beds are built by the aggregation of free living marine benthic coralline algae. Herein, we described phytobenthic communities associated with subtidal rhodolith beds in northeastern Brazil and tested the hypothesis that depth affects their structure. We compared macroalgal assemblages from depths of 10, 15 and 20 m. The genus Lithothamnion was dominant in these beds. Rhodolith density was similar at different depths, but volume decreases as depth increases. Sixty-seven species of fleshy algae were collected. The red algal order Ceramiales was dominant. A distinct community corresponds to each sampled depth. The shallower depth presented higher values for biomass, number of species, Shannon-Wiener diversity, and Pielou's evenness. When depth and water transparency increased, the number of species and the abundance of macroalgae decreased.     

Distribution of introduced Japanese macroalgae Undaria pinnatifida,Sargassum muticum (Phaeophyta) and Antithamnion Pectinatum (Rhodophyta) in the Lagoon of Venice

The present distribution of invasive seaweeds Undaria pinnatifida (Phaeophyta, Laminariales) and Sargassum muticum (Phaeophyta, Fucales) in the Lagoon of Venice is updated in this work. The distribution of Antithamnion pectinatum (Rhodophyta, Ceramiales) is presented for the first time. All these species have quickly colonized the hard substrata competing with indigenous species. Competition acts differently in the two species, U. pinnatifida competing for the substratum and S. muticum for light. On tidal flats composed of silt and sand, the growth of two brown seaweeds is restricted owing to the lack of a solid substrata. While Undaria pinnatifida and Sargassum muticum have colonized only some areas of the lagoon in a short vertical range (from 0 to -3 m), Antithamnion pectinatum is present on the three portmouths, along the inner shores of the city of Venice, and in the lagoon islands with extensive vertical distribution (from +0.2 to -8 m). This revised version was published online in September 2006 with corrections to the Cover Date.     

A mitochondrial marker for red algal intraspecific relationships

Intraspecific studies of red algae have relied on nuclear or plastid markers rather than mitochondrial data to address questions of systematics, biogeography or population genetics. In this study, primers were developed that spanned the noncoding intergenic region between the mitochondrial cytochrome oxidase subunit 2 and cytochrome oxidase subunit 3 genes. These primers were demonstrated to be successful on a variety of red algae in different orders: Gracilariales, Bonnemaisoniales and Ceramiales (families: Delesseriaceae, Ceramiaceae and Rhodomelaceae). Amplification products were between 450 and 320 bp in length, with variation in length shown among geographically distant isolates within a species. The region was variable within a single species, as shown for Bostrychia moritziana and B. radicans, and within populations of Caloglossa leprieurii. In the latter species, four mitochondrial haplotypes were observed in isolates from a single locality in Woolooware Bay, New South Wales, Australia. Analysis of hybrids between different mitochondrial haplotypes of B. moritziana revealed that the mitochondria are maternally inherited in this species. This is the first report of a mitochondrial marker that is variable within red algal populations and may lead to a better understanding of the population ecology of these important marine organisms.     

The benefits of polyandry in the free-spawning polychaete Galeolaria caespitosa

In many species, females are thought to benefit from polyandry due to the reduced risks of fertilization by genetically incompatible sperm. However, few studies that have reported such benefits have directly attributed variation in female reproductive success to the interacting effects of males and females at fertilization. In this paper, we determine whether male x female interactions influence fertilization in vitro in the free-spawning, sessile polychaete Galeolaria caespitosa. Furthermore, we determined whether polyandry results in direct fertilization benefits for females by experimentally manipulating the number of males contributing towards staged spawning events. To test for male x female interaction effects we performed an initial experiment that crossed seven males with six females (in all 42 combinations), enabling us to assess fertilization rates for each specific male-female pairing and attribute variation in fertilization success to males, females and their interaction. This initial experiment revealed a strong interaction between males and females at fertilization, confirming that certain male-female combinations were more compatible than others. A second experiment tested the hypothesis that polyandry enhances female reproductive success by exposing each female's eggs to either a single male's sperm (monandry) or the sperm from three males simultaneously (polyandry). We performed this second experiment at two ecologically relevant sperm concentrations. This latter experiment revealed a strong fertilization benefit of polyandry, independent of the effects of sperm concentration (which were also significant). We suggest that these direct fertilization gains arising from polyandry will constitute an important source of selection on females to mate multiply in nature.     

Latitudinal variation in intertidal algal community structure: the influence of grazing and vegetative propagation

Summary The hypothesis that sea urchin grazing and interactions with turf-forming red algae prevent large brown algae from forming an extensive canopy in the low intertidal zone of southern California was tested with field experiments at two study sites. Experimental removal of sea urchins resulted in rapid algal recruitment. Crustose coralline algae which typically dominate the substratum in areas with dense urchin populations were quickly overgrown by several species of short-lived green, brown and red algae. The removal of urchins also significantly increased the recruitment of two long-lived species of large brown algae (Egregia laevigata and Cystoseira osmundacea at one study site and E. laevigata and Halidrys dioica at the other). The experimental plots at both sites were eventually dominated by perennial red algae.A two-factorial experiment demonstrated that sea urchin grazing and preemption of space by red algae in areas where urchins are less abundant are responsible for the rarity of large brown algae in the low intertidal of southern California. The three dominant perennial red algae, Gigartina canaliculata, Laurencia pacifica and Gastroclonium coulteri, recruit seasonally from settled spores but can rapidly fill open space with vigorous vegetative growth throughout the year. These species encroach laterally into space created by the deaths of large brown algae or by other disturbances. Once extensive turfs of these red algae are established further invasion is inhibited. This interaction of algae which proliferate vegetatively with algae which recruit only from settled spores is analogous to those which occur between solitary and colonial marine invertebrates and between solitary and cloning terrestrial plants.It is suggested that a north-south gradient in the abundance of vegetatively propagating species, in grazing intensity and in the frequency of space-clearing disturbances, may account for latitudinal variation in intertidal algal community structure along the Pacific coast of North America.     

A new species of Procamallanus (Nematoda: Camallanidae) from Pacific eels (Anguilla spp.)

A new species of parasitic nematode, Procamallanus (Procamallanus) pacificus n. sp., is described from the stomach of the Pacific shortfinned eel, Anguilla obscura (type host), and from the speckled longfin eel, Anguilla reinhardtii, from northern New Caledonia (Melanesia, South Pacific); from Anguilla sp. (cf. obscura) from the Fiji Islands (Melanesia, South Pacific); and from the giant mottled eel Anguilla marmorata from Futuna Island (Wallis and Futuna Islands, Polynesia). Although a total of 450 nematodes were collected, all specimens were females; this suggests either an extremely rare occurrence of males or parthenogenetic reproduction in this species. Procamallanus pacificus differs markedly from all congeners from fish hosts in possessing a greater number (4-9) of caudal mucrons in the female and by other morphological features. This parasite might become a serious pathogen of cultured eels in the region of the South Pacific. Batrachocamallanus Jackson and Tinsley, 1995 is considered a junior synonym of Procamallanus Baylis, 1923, to which 2 species are transferred as Procamallanus occidentalis (Jackson and Tinsley, 1995) n. comb. and Procamallanus siluranae (Jackson and Tinsley, 1995) n. comb. One third-stage larva of Procamallanus (Spirocamallanus) sp. was also recorded from Anguilla sp. (cf. obscura) from the Fiji Islands.     

SPECIES OF ANTITHAMNION (RHODOPHYCEAE,CERAMIACEAE) OCCURRING ON THE SOUTHEAST AFRICAN COAST (NATAL)1

Three species of Antithamnion Naegeli were previously known from Natal, all reported since 1984. A more thorough collection of the flora in recent months has revealed that there are nine species in Natal, four of which are newly described. Vegetative characteristics are emphasized in this study of the taxonomy of Antithamnion, the most important being (1) size of axial cells, (2) form of whorl-branchlets, (3) size of whorl-branchlet basal cells, (4) size and shape of whorl-branchlet terminal cells, (5) size and position of gland cells, (6) primary or secondary derivation of indeterminate lateral branches, and (7) the relationship of primary indeterminate branches, if present, to normal arrangement of whorl-branchlets. Two closely related species, A. diminuatum Wollaston and A. eliseae sp. nov. have been grown in laboratory culture allowing for testing of stability and reliability of vegetative characteristics for use in taxonomic studies. Conclusions are that these criteria can be effectively used for these purposes. In addition to A. eliseae, A. nematocladellum, A. pterocladellum and A. adenocladellum also are described as new species. A tropical species previously known from the Caribbean and Red Seas, A. lherminieri Nasr, a species previously recorded from Japan, A. secundum Itono, a species previously known from California and Japan, A hubbsii Dawson, and two species from Australia, A. diminuatum Wollaston and A. divergens (J. Agardh) J. Agardh, are also described from Natal.     

The plastid genome of the red alga Laurencia

We present the 174,935 nt long plastid genome of the red alga Laurencia sp. JFC0032. It is the third plastid genome characterized for the largest order of red algae (Ceramiales). The circular‐mapping plastid genome is small compared to most florideophyte red algae, and our comparisons show a trend toward smaller plastid genome sizes in the family Rhodomelaceae, independent from a similar trend in Cyanidiophyceae. The Laurencia genome is densely packed with 200 annotated protein‐coding genes (188 widely conserved, 3 open reading frames shared with other red algae and 9 hypothetical coding regions). It has 29 tRNAs, a single‐copy ribosomal RNA cistron, a tmRNA, and the RNase P RNA.     

Geographic variation in the damselfish-red alga cultivation mutualism in the Indo-West Pacific

Background  

On coral reefs, damselfish defend their territories from invading herbivores and maintain algal turfs, from which they harvest filamentous algae. In southern Japan, intensive weeding of indigestible algae by Stegastes nigricans results in overgrowth by one filamentous alga, Polysiphonia sp. 1. Because this alga is highly susceptible to grazing and is competitively inferior to other algae, it survives only within the protective territories of this fish species, suggesting an obligate mutualism between damselfish and their cultivated alga. The wide distribution of damselfish species through the Indo-Central Pacific raises the question of whether this species-specific mutualism is maintained throughout the geographic range of the fish. To address this question, from all 18 damselfish species we conducted comprehensive surveys of algal flora within their territories throughout the Indo-West Pacific, and identified species of Polysiphonia using morphological examination and gene sequencing data.     

Red algal exotics on North Sea coasts

A total of ten red seaweed species are recognized as introduced into the North Sea from other parts of the world. These areAsparagopsis armata andBonnemaisonia hamifera (Bonnemaisoniales),Grateloupia doryphora (Halymeniales),Antithamnionella spirographidis, Antithamnionella ternifolia, Anotrichium furcellatum, Dasya baillouviana, ?Dasysiphonia sp.,Polysiphonia harveyi andPolysiphonia senticulosa (Ceramiales). The oldest of these isB. hamifera, introduced prior to 1890, while the most recent,?Dasysiphonia sp., was first found in 1994 and still requires taxonomic investigation. A variety of distribution patterns is seen, with geographical ranges varying from general within the North Sea to very restricted. The diversity of introduced red algae on eastern coasts of the North Sea is much greater than in the west. The most likely explanation for this pattern is that French coasts were the initial site of introduction for many of the seaweeds, which were then distributed northwards by the residual surface currents. Their increasing success in the Netherlands has probably been promoted by the drastically changed local hydrodynamic conditions which have also permitted the recent introduction of many native European species. Of the biological features of species that may favour their success as introductions, clonal vegetative propagation, often with specialized propagules or fragmentation mechanisms, is almost ubiquitous. Low-temperature tolerances can be inferred, but data are sparse. Many of the alien red algae in the North Sea contain anti-grazing compounds such as bromophenols, which may contribute to their invasive potential by deterring grazing sufficiently to permit establishment of an inoculum.     

Developmental morphology of Odonthalia floccosa (Ceramiales, Rhodophyta) in laboratory culture

Material of the red alga Odonthalia floccosa (Esper) Fal-kenberg (Rhodomelaceae, Ceramiales), collected from California, was cultured in the laboratory and its life-history was completed. Tetraspores grew into bipolar sporelings that differentiated into a colorless rhizoidal portion and a pigmented upright shoot. The sporelings became compressed apically and formed lateral branches in a regularly distichous manner that were congenitally fused with the main axis. These tetraspore germlings grew into diecious gametophytes. Male ga-metophytes produced numerous spermatangia on modified fertile branchlets (male trichoblasts) that possessed three to four monosiphonous, proximal segments. Female gametophytes formed a single pro-carp on the suprabasal segment of unbranched female trichoblasts. Cystocarps developed on the female gametophytes cocultured with male gametophytes and released viable carpospores that developed into fertile te-trasporophytes. Tetrasporangia were produced from the third and fourth periaxial cells in each of 12–45 successive fertile segments and provided three (two lateral and one basal) cover cells. The occurrence of both spermatangia and procarps on fertile trichoblasts in O. floccosa suggests that the alga is the most derived in these two characters among the species of the genus Odonthalia. This species is distributed in cold temperate regions in the North Pacific, and it should be excluded from the North Atlantic marine algal flora.     

Phylogenomics and multigene phylogenies decipher two new cryptic marine algae from California,Gelidium gabrielsonii and G. kathyanniae (Gelidiales,Rhodophyta)

Molecular surveys are leading to the discovery of many new cryptic species of marine algae. This is particularly true for red algal intertidal species, which exhibit a high degree of morphological convergence. DNA sequencing of recent collections of Gelidium along the coast of California, USA, identified two morphologically similar entities that differed in DNA sequence from existing species. To characterize the two new species of Gelidium and to determine their evolutionary relationships to other known taxa, phylogenomic, multigene analyses, and morphological observations were performed. Three complete mitogenomes and five plastid genomes were deciphered, including those from the new species candidates and the type materials of two closely related congeners. The mitogenomes contained 45 genes and had similar lengths (24,963–24,964 bp). The plastid genomes contained 232 genes and were roughly similar in size (175,499–177,099 bp). The organellar genomes showed a high level of gene synteny. The two Gelidium species are diminutive, turf‐forming, and superficially resemble several long established species from the Pacific Ocean. The phylogenomic analysis, multigene phylogeny, and morphological evidence confirms the recognition and naming of two new species, describe herein as G. gabrielsonii and G. kathyanniae. On the basis of the monophyly of G. coulteri, G. gabrielsonii, G. galapagense, and G. kathyanniae, we suggest that this lineage likely evolved in California. Organellar genomes provide a powerful tool for discovering cryptic intertidal species and they continue to improve our understanding of the evolutionary biology of red algae and the systematics of the Gelidiales.