Climate change and disease: bleaching of a chemically defended seaweed

Disease is emerging as an important impact of global climate change, due to the effects of environmental change on host organisms and their pathogens. Climate‐mediated disease can have severe consequences in natural systems, particularly when ecosystem engineers, such as habitat‐formers or top predators are affected, as any impacts can cascade throughout entire food webs. In temperate marine ecosystems, seaweeds are the dominant habitat‐formers on rocky reefs. We investigated a putative bleaching disease affecting Delisea pulchra, a chemically defended seaweed that occurs within a global warming ‘hot‐spot’ and assessed how patterns of this phenomenon were influenced by ocean temperature, solar radiation, algal chemical defences and microbial pathogens. Warmer waters were consistently and positively correlated with higher frequencies of bleaching in seaweed populations, but patterns of bleaching were not consistently influenced by light levels. Bleached thalli had low levels of antibacterial chemical defences relative to healthy conspecifics and this was observed across entire thalli of partially bleached algae. Microbial communities associated with bleached algae were distinct from those on the surfaces of healthy seaweeds. Direct testing of the importance of algal chemical defences, done here for the first time in the field, demonstrated that they protected the seaweed from bleaching. Treatment of algal thalli with antibiotics reduced the severity of bleaching in experimental algae, especially at high water temperatures. These results indicate that bleaching in D. pulchra is the result of temperature‐mediated bacterial infections and highlight the potential for warming to influence disease dynamics by stressing hosts. Understanding the complex ways in which global change may affect important organisms such as habitat‐forming seaweeds, is essential for the management and conservation of natural resources.     

European bioconversion projects and realizations for macroalgal biomass : Saint-Cast-Le-Guildo (France) experiment

Proliferation of macroalgae is a world-wide problem with 50,000 m³ of drift Ulva harvested per year in Brittany and about 1.0 to 1.2 million tons growing in the Venice lagoon. This biomass may be treated by bioconversion (aerobic or anaerobic fermentation) to give useful products (gas, fertilizers or others) and to remove a source of environmental pollution. Such a treatment also may be applied to cultivated or harvested seaweds and to seaweed industry residues.Studies of seaweed methanization showed Laminaria an especially good substrate and Ulva a possible substrate. Research led to a defined way of treating drift algae, encompassing natural hydrolysis and pressing with methanization of the juices.The most advanced full-scale realization for algal biomass utilization is the C.A.T.-Quatre-Vaulx composting plant in Saint-Cast-Le-Guildo (Brittany, France). It produced from seaweed, wood and animal dung a biological quality compost that is competitive with the traditional market products.     

The role of Ulva spp. as a temporary nutrient sink in a coastal lagoon with oyster cultivation and upwelling influence

Bahía San Quintín is a coastal lagoon with large Ulva biomass and upwelling influence. Previous observations suggest that Ulva has increased recently as a result of oyster cultivation. To evaluate the possible role of Ulva as a temporary nutrient sink, biomass and tissue C, N, and P were determined seasonally. Maximum biomass was present during spring and early summer (1,413–1,160 t (dry)) covering about 400 ha. However, the biomass decreased to 35 t (dry) by winter. The mean annual percentage of Ulva C, N, and P was 28%, 2%, and 0.14%, respectively. This study shows that Ulva can store up to 28 t of N and 2 t of P in Bahía San Quintín during spring–summer. Ulva may be displacing the seagrass Zostera marina subtidal beds. A partial removal of the seaweed would reduce the risk of eutrophication and the displacement of eelgrass beds.     

Physical effects of habitat‐forming species override latitudinal trends in temperature

Latitudinal and elevational temperature gradients (LTG and ETG) play central roles in biogeographical theory, underpinning predictions of large‐scale patterns in organismal thermal stress, species' ranges and distributional responses to climate change. Yet an enormous fraction of Earth's taxa live exclusively in habitats where foundation species modify temperatures. We examine little‐explored implications of this widespread trend using a classic model system for understanding heat stresses – rocky intertidal shores. Through integrated field measurements and laboratory trials, we demonstrate that thermal buffering by centimetre‐thick mussel and seaweed beds eliminates differences in stress‐inducing high temperatures and associated mortality risk that would otherwise arise over 14° of latitude and ~ 1 m of shore elevation. These results reveal the extent to which physical effects of habitat‐formers can overwhelm broad‐scale thermal trends, suggesting a need to re‐evaluate climate change predictions for many species. Notably, inhabitant populations may exhibit deceptive resilience to warming until refuge‐forming taxa become imperiled.     

Bioactive compounds in seaweed: functional food applications and legislation

Seaweed is more than the wrap that keeps rice together in sushi. Seaweed biomass is already used for a wide range of other products in food, including stabilising agents. Biorefineries with seaweed as feedstock are attracting worldwide interest and include low-volume, high value-added products and vice versa. Scientific research on bioactive compounds in seaweed usually takes place on just a few species and compounds. This paper reviews worldwide research on bioactive compounds, mainly of nine genera or species of seaweed, which are also available in European temperate Atlantic waters, i.e. Laminaria sp., Fucus sp., Ascophyllum nodosum, Chondrus crispus, Porphyra sp., Ulva sp., Sargassum sp., Gracilaria sp. and Palmaria palmata. In addition, Undaria pinnatifida is included in this review as this is globally one of the most commonly produced, investigated and available species. Fewer examples of other species abundant worldwide have also been included. This review will supply fundamental information for biorefineries in Atlantic Europe using seaweed as feedstock. Preliminary selection of one or several candidate seaweed species will be possible based on the summary tables and previous research described in this review. This applies either to the choice of high value-added bioactive products to be exploited in an available species or to the choice of seaweed species when a bioactive compound is desired. Data are presented in tables with species, effect and test organism (if present) with examples of uses to enhance comparisons. In addition, scientific experiments performed on seaweed used as animal feed are presented, and EU, US and Japanese legislation on functional foods is reviewed.     

Acclimation of bloom‐forming and perennial seaweeds to elevated pCO2 conserved across levels of environmental complexity

Macroalgae contribute approximately 15% of the primary productivity in coastal marine ecosystems, fix up to 27.4 Tg of carbon per year, and provide important structural components for life in coastal waters. Despite this ecological and commercial importance, direct measurements and comparisons of the short‐term responses to elevated pCO₂ in seaweeds with different life‐history strategies are scarce. Here, we cultured several seaweed species (bloom forming/nonbloom forming/perennial/annual) in the laboratory, in tanks in an indoor mesocosm facility, and in coastal mesocosms under pCO₂ levels ranging from 400 to 2,000 μatm. We find that, across all scales of the experimental setup, ephemeral species of the genus Ulva increase their photosynthesis and growth rates in response to elevated pCO₂ the most, whereas longer‐lived perennial species show a smaller increase or a decrease. These differences in short‐term growth and photosynthesis rates are likely to give bloom‐forming green seaweeds a competitive advantage in mixed communities, and our results thus suggest that coastal seaweed assemblages in eutrophic waters may undergo an initial shift toward communities dominated by bloom‐forming, short‐lived seaweeds.     

Changes in biomass and botanical composition of beach-cast seaweeds in a disturbed coastal area from Argentine Patagonia

Trends in wrack composition and biomass, and its relationship with the anthropogenic impact were studied along a coastal area in Nuevo Gulf (south Patagonia) in front of Puerto Madryn city. Beach-cast macroalgae composition was sampled from 1992 to 1999 in the Puerto Madryn beaches and in several other nearby beaches in 1993, 1996 and 1998. Historical information was based on local knowledge and observations reported by marine biologists who worked in the area. The botanical composition of the beach-cast macroalgae in Puerto Madryn indicates a succession in the dominance from Codium spp. to Ulva spp. during the 1990s and from Ulva to Undaria pinnatifida since 1998, accompanied by a significant decrease in biomass of Gracilaria gracilis and Macrocystis pyrifera. The increase of the opportunist species such as Ulva may be supported by the continuous delivery of waste waters into the Nuevo Gulf while the dominance of U. pinnatifida may be associated with port activities. During the sampling period the highest wrack biomass values were recorded in spring and summer. The beach-cast seaweed biomass harvested by the municipality of Puerto Madryn during beach cleaning operation, ranged between 2500 and 12000 t year^–1 ( 200 and 960 t dry weight). Wrack harvesting produces an environmental impact by removing sand from the beach and affecting coastal communities. Composting of wrack is proposed as one of the environmental alternatives to land disposal.     

Management of natural <Emphasis Type="Italic">Ulva</Emphasis> spp. blooms in San Quintin Bay,Baja California: Is it justified?

According to Zertuche-González et al. (2009), Ulva spp. blooms, favored by oyster cultivation, are likely displacing subtidal meadows of Zostera marina in San Quintin Bay, Baja California. The authors propose a partial removal of the seaweed to reduce the risk of eutrophication and eelgrass displacement in the bay. We warn about the removal of Ulva spp. biomass by raising six arguments that emphasize the necessity of a historical and ecosystem-based management for San Quintin Bay. First, processes other than competitive exclusion by Ulva spp. blooms more likely explain changes of Z. marina subtidal meadows in the past decade. Second, there is no consistent evidence that oyster cultivation is promoting blooms of Ulva spp. and the loss of eelgrass. Third, the removal of Ulva spp. biomass is based on experiences of heavily anthropogenically eutrophic systems, while San Quintin Bay is not. Fourth, the proposed course of actions to restore eelgrass meadows ignores general historical baselines of estuarine and coastal systems by confusing what it means to be “pristine.” Fifth, despite the important experimental evidence indicating strong top–down control in temperate seagrass meadows, Zertuche-González et al. (2009) underestimated the capacity of consumers in structuring dynamics of vegetated soft-bottom communities in San Quintin Bay. Sixth, Ulva expansa may exert positive effects on seagrass ecosystem properties and functions. Instead, we propose protection against the propagation of unsustainable practices in the bay, and the reintroduction of large consumers that are now absent in this ecosystem. An ecosystem-based analysis of the role of Ulva spp. on eelgrass dynamics is needed.     

SEASONALITY OF SELECTED NUTRITIONAL CONSTITUENTS OF EDIBLE DELMARVA SEAWEEDS

Edible seaweeds have not been thoroughly explored for food, medicinal, or industrial purposes in the United States. This study compared selected proximate constituents of three edible seaweed species (Ulva lactuca L., Fucus vesiculosus L., and Gracilaria tikvahiae McLachan) at two sites for possible future development as a food crop on the Delmarva Peninsula. Sampling was conducted bimonthly at Chincoteague Memorial Park, Virginia, and Indian River Inlet, Delaware, from 2005 to 2008. Proximate constituents of moisture, ash, dietary fiber, proteins, and fat were measured seasonally and calorific values were calculated. Data were analyzed using correlation, paired samples t‐tests and one‐ and two‐way ANOVA. Significant variations in the proximate constituents were found among seasons, species, and between sites. The brown seaweed (Fucus) at both sites had higher fiber, fat, and ash (mineral) content than the green (Ulva) or the red (Gracilaria). Ulva and Gracilaria had higher protein content than Fucus. Seaweeds from Delaware had more fat, ash, and protein than from Virginia, potentially because of the more polluted, nutrient rich environment at the Delaware site. Positive correlations between seaweed fat and protein content may indicate an increase in the synthesis of both components under optimal growth conditions. Species' physiology differences and the water quality at the two sites likely impacted proximate constituent values. This study contributed new information to the existing body of knowledge in the areas of nutrition and ecology of seaweeds and their potential as a cash crop.     

Ulva diversity in the Yellow Sea during the large‐scale green algal blooms in 2008–2009

In the Yellow Sea of China, large‐scale green tides have broken out for three consecutive years from 2007 to 2009. As part of the efforts to localize the algal source, two cruises were conducted in the early stage and the outbreak stage of the bloom in 2009. We analyzed the morphological and genetic diversity of drifting Ulva specimens and culture‐derived isolates from seawater sampled in different localities. For phylogenetic analyses, the nuclear encoded ribosomal DNA internal transcribed spacer region (ITS nrDNA) and the plastid encoded large subunit of ribulose‐1, 5‐bisphosphate carboxylase/oxgenase gene (rbcL) were used. Our molecular and morphological data indicate that the dominant free‐floating Ulva species in 2008 and 2009 possibly belonged to a single strain of the U. linza‐procera‐prolifera (LPP) clade. The ITS sequences from bloom‐forming algal samples with dense branches were identical to those from U. linza‐like specimens without branches derived from the Yellow Sea. Microscopic individuals of the dominant Ulva strain were detected in eight stations, revealing that spore dispersal in the water helped to enlarge biomass in the water during the outbreak stage of green tide in the Yellow Sea.     

Ulva: An emerging green seaweed model for systems biology

Green seaweeds exhibit a wide range of morphologies and occupy various ecological niches, spanning from freshwater to marine and terrestrial habitats. These organisms, which predominantly belong to the class Ulvophyceae, showcase a remarkable instance of parallel evolution toward complex multicellularity and macroscopic thalli in the Viridiplantae lineage. Within the green seaweeds, several Ulva species (“sea lettuce”) are model organisms for studying carbon assimilation, interactions with bacteria, life cycle progression, and morphogenesis. Ulva species are also notorious for their fast growth and capacity to dominate nutrient-rich, anthropogenically disturbed coastal ecosystems during “green tide” blooms. From an economic perspective, Ulva has garnered increasing attention as a promising feedstock for the production of food, feed, and biobased products, also as a means of removing excess nutrients from the environment. We propose that Ulva is poised to further develop as a model in green seaweed research. In this perspective, we focus explicitly on Ulva mutabilis/compressa as a model species and highlight the molecular data and tools that are currently available or in development. We discuss several areas that will benefit from future research or where exciting new developments have been reported in other Ulva species.     

Growing Ulva (Chlorophyta) in integrated systems as a commercial crop for abalone feed in South Africa: a SWOT analysis

Over 1,000t fresh weight of Ulva was cultivated on South African abalone farms in 2007, primarily for feed, but in one case to allow partial re-circulation by nutrient removal. The potential of this new commercial crop is analysed. Material is collected from natural free-floating populations and at least four species are commonly grown, with different ecophysiological characteristics. A brown epiphyte, Myrionema strangulans, causes a disease of the Ulva, which is currently managed by farmers by re-stocking. The main potential threat is that some farmers are wary of integrated systems, fearing they may promote disease in abalone, although Ulva has been grown in abalone effluent and fed back to the abalone on one farm for 6 years without adverse effects. Opportunities exist for the expansion of Ulva cultivation via further spread of the abalone industry, the inclusion of seaweed raceways in proposed fish-farming activities, or the potential for the inclusion of high quality cultivated Ulva as a constituent in aquafeed. A conservative estimate of production over a full year from these raceway systems was 26.1 g dw m⁻² day⁻¹ (2006) and 19.7 g dw m⁻² day⁻¹ (2007), similar to maximum figures for total annual microalgal biomass production in outdoor systems.     

Differential toxic effects of Ulva lactuca (Chlorophyta) on the herbivorous gastropods,Littorina littorea and L. obtusata (Mollusca)

Members of the genus Ulva are widespread and abundant in intertidal and shallow subtidal areas but there are conflicting data regarding susceptibility to herbivory. While some studies have documented that Ulva spp. were favored by a diversity of marine herbivores, other work has revealed herbivore deterrence. We investigated grazing and growth rates of the littorinid species, Littorina littorea and L. obtusata, when offered Fucus vesiculosus, Ascophyllum nodosum, Ulva lactuca, and Chondrus crispus, highlighting distinctive vulnerabilities to toxic effects of U. lactuca. Ulva lactuca was the preferred food of L. littorea, while L. obtusata showed no grazing on this ephemeral algal species. In contrast, F. vesiculosus was highly preferred by L. obtusata. Although L. littorea demonstrated a grazing preference for U. lactuca, growth rate of this gastropod species was nearly 3× greater when fed F. vesiculosus, suggesting a non‐lethal, negative effect of U. lactuca on L. littorea with long‐term exposure. Mortality of L. obtusata ranged from 0% to 100% when held in the presence of various Ulva densities for 1 week, and Ulva exudate depressed herbivory of this gastropod. We conclude that the water‐soluble, toxic exudate produced by U. lactuca in response to herbivory had allelochemical properties, and may contain a cleavage product (acrylic acid) of dimethylsulfoniopropionate or reactive oxygen species (i.e., H₂O₂). Observed differences in susceptibility to Ulva toxicity by the littorinid species may be related to generalist versus specialist feeding and habitat strategies.     

Interaction of Mercuric Ions with Different Marine Algal Species

The use of different seaweeds such as Sargassum sp., Turbinaria conoides, and Ulva sp. in removing mercury(II) from aqueous solutions were investigated. The initial experimental runs, conducted at different equilibrium pH conditions, demonstrated that brown seaweeds outperformed green seaweed in Hg(II) biosorption at all pH conditions. In particular, at pH 5, maximum biosorption capacities of 170.3 and 145.8 mg/g were recorded for the brown seaweeds T. conoides and Sargassum sp., respectively, compared with 138.4 mg/g for the green seaweed Ulva sp. Isotherm data were modeled and interpreted using the Langmuir, Freundlich, Redlich-Peterson, and Toth models, with the latter described the Hg(II) isotherms with high correlation coefficients and low % error values. The kinetic data indicate the rapidity of the biosorption process, with the equilibrium achieved within 90 min. Several models, including the Elovich, pseudo-first-order, and pseudo-second-order models, were examined for their suitability with the present data; the correlation coefficient and % error values, along with better prediction of equilibrium uptake values, favored the pseudo-first-order model. The desorption experiments were highly successful for T. conoides biomass with 0.05 M HCl, whereas for the other two seaweeds, 0.05 M HCl resulted in high biomass weight loss. Reusing T. conoides biomass in three successive sorption-desorption cycles resulted in only 8.8% reduction in Hg(II) biosorption capacity compared with its original uptake.     

Freshwater population of Ulva flexuosa (Ulvaceae,Chlorophyta) as a food source for great pond snail: Lymnaea stagnalis (Mollusca,Lymnaeidae)

Cosmopolitan species of the genus Ulva (Ulvaceae; Chlorophyta) that populate the littoral zone of marine habitats constitute a staple diet for a variety of organisms, particularly snails, shellfish, polychaetes, and birds. Occurrence of Ulva species (e.g., U. flexuosa and U . prolifera) has also been observed in freshwater inland ecosystems that have no contact with saline water. However, the influence of the development of macroalgal mats of Ulva on indigenous organisms in limnic ecosystems has not been established. This study investigates the trophic relationships between Ulva flexuosa and one species of snail from freshwater habitats in central Europe. During the summer, the great pond snail (Lymnaea stagnalis) consumed Ulva as a source of nutrition even when other algae and plants were available. Lymnaea stagnalis consumed an average of 100 mg of Ulva thalli per day. This level of biomass exceeded the consumption of an alternative food source, the shoots of Elodea canadensis. Ulva thalli are more actively consumed by great pond snails than Elodea shoots, and this is expressed in terms of the differences of biomass consumption. It was also observed that the interior of the monostromatic tubular thalli of Ulva flexuosa serves as a protective shelter for juvenile great pond snails.     

Vertical Distribution, Size Structure, and Habitat Associations of Four Blenniidae Species on Gas Platforms in the Northcentral Gulf of Mexico

We counted individuals of the family Blenniidae and estimated their sizes on gas platforms southeast of Dauphin Island, Alabama. We observed species abundance decreasing as depth increased. Fish sizes also decreased with depth. The most abundant species was molly miller, Scartella cristata, followed by plumed blenny, Hypleurochilus multifilis, tessellated blenny, Hypsoblennius invemar, and seaweed blenny, Parablennius marmoreus. Total blenny abundance was positively related to the barnacle, Megabalanus antillensis, and dissolved oxygen concentrations, and inversely related to Anthozoa. Individually, S. cristata was correlated with M. antillensis, and inversely related to salinity, while the other blenny species showed more complex correlations to invertebrates. As a community, blennies showed a clear separation based on depth independent of offshore/inshore sites and sample date based on multidimensional scaling analyses. Our study suggests that attached invertebrates, particularly M. antillensis, provided a predation refuge, allowing these blenny species to exist in an otherwise unsuitable habitat, i.e. open shallow waters of the northern Gulf of Mexico.     

A Molecular Survey of Ulva (Chlorophyta) in Temperate Australia Reveals Enhanced Levels of Cosmopolitanism

The green algal genus Ulva includes a speciose group of marine macroalgae inhabiting shallow seas worldwide. Although algal blooms in Asia highlight the opportunistic nature of several “nuisance” species, recent research clearly reveals important positive benefits of Ulva. Applied research requires accurate, reliable, and rapid identification, however, identification of Ulva spp. has met with con‐siderable difficulty. Consequently, many have turned to molecular markers to aid in taxonomy. Previous studies of plants and algae have relied heavily on ITS and rbcL. Recently, tufA has been presented as a suitable barcoding gene to facilitate species‐level identification of green macroalgae and it is used here to explore the diversity of Ulva spp. in temperate Australia. Ninety Ulva specimens collected from 38 sites across five states were sequenced for this gene region with exemplars from each genetic group also sequenced for rbcL to test for congruence. Collections of Australian Ulva spp. were compared to samples from Asia and North America and exhibited trends consistent with recent studies in terms of species relationships. Results support an overwhelmingly cosmopolitan flora in temperate Australia that contrasts with other Australasian surveys of Ulva that report a greater number of endemics and new species. Four new records, as well as numerous range extensions for taxa already known from the country, are documented. Evidence for three nonindigenous Ulva species in temperate Australia is discussed.     

Trophic cascades in a temperate seagrass community

We assessed the relative importance of bottom–up and top–down processes in structuring an eelgrass community in Sweden, a system impacted both by eutrophication and overfishing. Using artificial seagrass as substrate, we manipulated nutrient levels and predator abundance in a full‐factorial cage‐experiment. The results revealed a seagrass community dominated by strong top–down processes controlling the aggregate biomass of mesograzers and macroalgae. In the absence of predators the large amphipod Gammarus locusta became very abundant resulting in a leaf community with low biomass of algae and smaller mobile fauna. One enclosed gobid fish predator reduced the abundance of adult G. locusta by >90%, causing a three to six times increase in the biomass of algae, smaller mesograzers and meiofauna. Numerous small predators in uncaged habitats reduced the biomass of G. locusta and other mesograzers by >95% in comparison to the fish treatment, further increasing the biomass of epiphytic algae and meiofauna. Although water column nutrient enrichment caused a temporal bloom of the filamentous macroalgae Ulva spp., no significant nutrient‐effects were found on the algal community at the end of the experiment. The only lasting nutrient‐effect was a significant increase in the biomass of G. locusta, but only in the absence of ambient predators. These results demonstrate that mesograzers can respond to enhanced food supply, increase their biomass and control the algal growth when predation rates are low. However, in the assessed system, high predation rates appear to make mesograzers functionally extinct, causing a community‐wide trophic cascade that promotes the growth of ephemeral algae. This top–down effect could penetrate down, despite a complex food‐web because the interaction strength in the community was strongly skewed towards two functionally dominant algal and grazer species that were vulnerable to consumption. These results indicate that overexploitation of gadoid fish may be linked to increased macroalgal blooms and loss of eelgrass in the area through a trophic cascade affecting the abundance of mesograzers.     

Ulva (Chlorophyta,Ulvales) Biodiversity in the North Adriatic Sea (Mediterranean,Italy): Cryptic Species and New Introductions

Ulva Linnaeus (Ulvophyceae, Ulvales) is a genus of green algae widespread in different aquatic environments. Members of this genus show a very simple morphology and a certain degree of phenotypic plasticity, heavily influenced by environmental conditions, making difficult the delineation of species by morphological features alone. Most studies dealing with Ulva biodiversity in Mediterranean waters have been based only on morphological characters and a modern taxonomic revision of this genus in the Mediterranean is not available. We report here the results of an investigation on the diversity of Ulva in the North Adriatic Sea based on molecular analyses. Collections from three areas, two of which subject to intense shipping traffic, were examined, as well as historical collections of Ulva stored in the Herbarium Patavinum of the University of Padova, Italy. Molecular analyses based on partial sequences of the rbcL and tufA genes revealed the presence of six different species, often with overlapping morphologies: U. californica Wille, U. flexuosa Wulfen, U. rigida C. Agardh, U. compressa Linnaeus, U. pertusa Kjellman, and one probable new taxon. U. californica is a new record for the Mediterranean and U. pertusa is a new record for the Adriatic. Partial sequences obtained from historical collections show that most of the old specimens are referable to U. rigida. No specimens referable to the two alien species were found among the old herbarium specimens. The results indicate that the number of introduced seaweed species and their impact on Mediterranean communities have been underestimated, due to the difficulties in species identification of morphologically simple taxa as Ulva.     

Molecular phylogenetic analyses of the Japanese Ulva and Enteromorpha (Ulvales, Ulvophyceae), with special reference to the free-floating Ulva

In order to elucidate the species composition of free‐floating Ulva that cause green tide in several bays in Japan, and to clarify the generic status of Ulva and Enteromorpha (Ulvales, Ulvophyceae), the nuclear encoded internal transcribed spacer (ITS) region including the 5.8S gene and the plastid encoded large subunit of ribulose‐1, 5‐bisphosphate carboxylase/ oxgenase (rbcL) gene sequences for 15 species were determined. Both ITS and rbcL analyses indicate that free‐floating Ulva samples are divided into four different lineages that correspond to Ulva lactuca Linnaeus, U. pertusa Kjellman, U. armoricana Dion etal. and U. fasciata Delile. These four species are distinguished by cell morphology including the arrangement of cells, the shape and size of cells and the position of chloroplasts. Molecular data also indicated that Ulva and Enteromorpha are not separated as respective monophyletic groups within a large monophyletic clade and congeneric as shown by previous molecular studies using the ITS sequences alone. This strongly suggests that these genera are congeneric and Enteromorpha should be reduced to the synonym of Ulva.