Growth and reproductive patterns of <Emphasis Type="Italic">Undaria pinnatifida</Emphasis> sporophytes in a cultivation farm in Busan,Korea

Monthly growth and reproduction of Undaria pinnatifida sporophytes were examined over a period of 5 months in a cultivation farm in Korea. A total of 11 characters of Undaria were measured to determine a reliable morphological character representing its growth and reproduction. Plant weight of Undaria sporophytes increased steadily over the experimental period, but it increased in four different ways. Undaria pinnatifida increased body weight by growth in length and width (October–early December), and by growth in width with the thickening of blade and stipe when sporophytes began to be fertile (December–January). In the middle of January, growth in length and width had almost stopped with the maturation of Undaria sporophytes. Finally, the weight of Undaria increased again by growth in width at the end of February. Present results indicate that Undaria sporophytes increase body weight by growth in length and width at different times, and the relationship between reproduction and vegetative growth is exclusive. Plant weight was positively correlated and fitted well with stipe width and blade width. The blade of Undaria was very thin (ca. 254 μm) and breakable by wave action, but its stipe was strong and relatively thick (ca. 8.7 mm). Furthermore, the fertility of U. pinnatifida was fitted better with stipe width than blade width. Thus, we suggest that the stipe width is the most feasible character with which to estimate the growth and reproduction of U. pinnatifida sporophytes in the cultivation farm.     

First Report of the Asian kelp Undaria pinnatifida in the Northeastern Pacific Ocean

The Asian kelp Undaria pinnatifida was discovered in southern California in the spring of 2000, and by the summer of 2001 had been collected at several California sites from Los Angeles to Long Beach Harbors and Catalina Island as far north as Monterey Harbor. From its native range in the northwestern Pacific, Undaria has now been introduced to the northeastern and southwestern Atlantic and the southwestern and northeastern Pacific, through a combination of intentional transport for cultivation, accidental transport with oysters, as fouling on vessel hulls, and possibly other means. In the northeastern Pacific, water temperatures are suitable for its establishment from at least Baja California to British Columbia, where it is likely to grow well in sheltered and partially sheltered waters.     

The Invasive Kelp Undaria Pinnatifida (Phaeophyceae, Laminariales) Reduces Native Seaweed Diversity in Nuevo Gulf (Patagonia, Argentina)

The kelp Undaria pinnatifida(Phaeophyceae) is a seaweed native to northeast Asia, but during the last two decades, it has been accidentally or intentionally introduced in several temperate coasts worldwide. In central Patagonia (Argentina), this species was first detected in late 1992, and it is progressively spreading from the point of introduction. Through a manipulative experiment involving Undaria removal in 2001, we found that its presence is associated with a dramatic decrease in species richness and diversity of native seaweeds in Nuevo Gulf. Future prospects are worrisome, as, in addition to the negative impact from a biodiversity viewpoint, native commercial macroalgae and invertebrates might also be affected.     

162 Interactions Between Planktonic Microalgae and Protozoan Grazers

Invasive algal species have the potential to change the structure and ecology of native algal communities. One well‐known invader, the large Japanese kelp Undaria pinnatifida, has recently become established at several locations along the central and southern California coast (Monterey, Santa Barbara, Catalina, and others). Previous to its introduction in the northeastern Pacific, Undaria has become established along the coastlines of several countries, including Australia, New Zealand, Argentina, England, and France. However, the seasonal population dynamics, rate of spread, and impact on local communities at each invasion site varies. Undaria in the Santa Barbara, CA harbor exhibits two distinct recruitment pulses per year (fall, late winter), with nonoverlapping generations of adult individuals. Individuals can grow rapidly and become reproductive a month after appearing as recruits (2–3 cm long), indicating a potential for rapid spread. However, Undaria may be effectively controlled by grazing via natural recruitment of the kelp crab Pugettia producta. However, Undaria invasions in other California invasions have not been controlled by herbivory, and Undaria populations in these areas have the potential to compete with a wide diversity of native California kelp species for habitat space and light.     

Undaria mitochondrial large subunit ribosomal DNA (Phaeophyceae, Alariaceae): Structural properties and use for phylogenetic analysis

We sequenced mitochondrial 23S ribosomal DNA (rDNA) from two Undaria species and compared them with homologous sequences from related brown algae. The 23S rDNA sequences from Undaria pinnatifida (Harvey) Suringar were 2707 bp in length, of which 1290–1292 sites were assigned to stem regions and the rest to loop regions in the rRNA secondary structure model. In Undaria undarioides (Yendo) Okamura, the sequences were 2708–2709 bp in length and 1290 sites were assigned to stems. There were three sequence types among the seven U. pinnatifida individuals examined and two among the four U. undarioides individuals. A maximum parsimony analysis of these sequence data with those from related brown algal species resolved a robust monphyly of each of the two Undaria species, suggesting the utilities of 23S rDNA for phylogeny at species or intraspecific level in brown algae. Accounting for variations among sequences parsimoniously, sequence changes were sorted into 423 events, of which 165 were mapped to stems of the secondary structure.     

Undaria pinnatifida (Laminariales,Phaeophyta) 12 years after its introduction into the Atlantic Ocean

Since its first introduction into the Atlantic for farming purposes in 1983, the brown alga Undaria pinnatifida has been found at several sites along the European coasts. The present study deals with an ecological experiment that was performed at the island of Ushant (Brittany, France) in order to check the ability of Undaria to recruit to a rocky substratum in situ. On the one hand, the authors were unsuccessful in getting Undaria to colonize a previously denuded area of shore, which was subsequently colonized by other native species, a result that may speak for a competition between Undaria and the local kelps. On the other hand, evidence is presented that Undaria pinnatifida has become a member of the Atlantic flora for good. The sporophytes can be found growing from low water mark of neap tide (+1.5 m) down to at least –15 m below lower water mark of spring tide. Its propagation is thought to occur in at least two ways: either step by step from cultivation sites along the coasts, or from the hulls of ships sailing from harbour to harbour. The preference of Undaria for settlement is on artificial structures.     

First Record of Undaria Pinnatifida (Laminariales, Phaeophyta) in Southern Patagonia, Argentina

The Japanese kelp Undaria pinnatifida was found on 7th September 2005 in Ría Deseado (Santa Cruz, Argentina). This invasive algae was registered from the lower intertidal to the upper levels of the subtidal zone. The sporophytes were fixed to slabs, gravel and rocks of variable sizes and were also present as epibionts of tunicates. The sporophytes total length ranged between 3 and 88 cm, with a mean of 28.75 cm in the intertidal and 38.3 cm in the subtidal. The density and biomass increased from the intertidal to the subtidal. The kelp population showed a higher number of small immature individuals in the intertidal than␣subtidal where the organisms were bigger and with more individual biomass. This first record of U. pinnatifida in Southern Patagonia extends its southern distribution limit in Argentina, showing progressive expansion in the southwestern Atlantic. More surveys must be conducted to understand the evolution of U. pinnatifida invasion and its impact on the native benthic community.     

An investigation of dispersal of the introduced brown alga Undaria pinnatifida (Harvey) Suringar and its competition with some species on the man-made structures of Torquay Marina (Devon, UK)

As part of a study into the ecology of the recently introduced macroalga Undaria pinnatifida in the British Isles, the competitive ability of this adventive kelp with native species was investigated in Torquay Marina (Devon, UK). Sites of introduction were surveyed for the presence of native species filling a similar ecological niche and long-term competition experiments set up over 6 years. Experiments involving clearance, removal and monitoring showed that Undaria has a low competitive ability in the infralittoral and sublittoral of a rocky breakwater. Undaria did not manage to colonise the nearby (200 m) shoreline for at least 7 years from the date of its original introduction to the floating pontoons of Torquay Marina. Undaria did out-compete native species in a marina environment: two kelp species, Laminaria digitata and Laminaria saccharina, and a sea squirt, Styela clava.     

The occurrence of pinhole disease in relation to developmental stage in cultivated <Emphasis Type="Italic">Undaria pinnatifida</Emphasis> (Harvey) Suringar (Phaeophyta) in Korea

Infection rates of pinhole disease in farmed Undaria pinnatifida (Harvey) Suringar were examined between November 2002 and April 2003 at an Undaria farm at Wando on the southwestern coast of Korea. The growth stages of U. pinnatifida can be defined as: a growth phase from December to the end of February, and a decay phase from March to April. There was a significant increase in the level of infection with pinhole disease (measured as the number of pinholes per cm²) between December and April (P < 0.01). Infection progressed from the upper part of the frond in December to the basal part in April. The infection rate of pinhole disease in relation to growth phase on farms over time is limited by the harvest of the seaweed crop at the end of the cultivation period in April. The midrib, sporophyll and stipe showed no signs of infection at any time during the cultivation period. The reduced physiological activity and tissue aging that occurred over time in the sporophytic life phase is the probable cause of the increasing infection rate of the disease.     

Cultivation of a hybrid of free-living gametophytes between Undariopsis peterseniana and Undaria pinnatifida: morphological aspects and cultivation period

The kelp Undariopsis peterseniana is warm-water-tolerant, and consequently, there is currently considerable interest in developing commercial cultivation techniques for this species in Korea. U. peterseniana plants have been successfully transferred to the northern coast of Korea beyond their original habitat in Jeju Island (33°30′08.65″N, 126°55′39.02″E). In this study, we cultured a hybrid kelp consisting of a cross between free-living gametophytes of U. peterseniana and U. pinnatifida in an attempt to extend the culture period of Undaria which is an important species for both the abalone industry and for commercial seaweed mariculture for human food applications. Morphological characters and cultivation period were compared between the parent thalli and the hybrid. The cultivation experiment was conducted in Wando, on the southern coast of Korea (34°26′18.68″N, 127°05′43.88″E). The morphological characteristics of the hybrid thalli were intermediate between the two species having shallow pinnated blades and a reduced reproductive organ. Hybrid thalli showed faster growth rates, 1.5 times greater biomass, and a longer cultivation period than the parent thalli. The hybrid strain possessed characteristics that indicate it could be used as an alternative kelp source to supply the abalone feed industry.     

The genetic analysis and germplasm identification of the gametophytes of Undaria pinnatifida (Phaeophyceae) with RAPD method

Eleven pairs of Undaria pinnatifida (Harv.) Suringar gametophytes were identified with random amplified polymorphic DNA (RAPD) technique. After screening 100 primers, 20 ten-base primers were determined for the RAPD analysis. A total of 312 polymorphic loci were obtained, of which 97.7% were polymorphic. The primer S198 was found to distinguish all the selected Undaria pinnatifida gametophytes. The genetic distances between each two of the twenty-two U. pinnatifida gametophytes ranged from 0.080 to 0.428, while the distances to the Laminaria was 0.497 on average. After reexamination, two sequences characterized amplification region (SCAR) markers were successfully converted, which could be applied to U. pinnatifida germplasm identification. All these results demonstrated the feasibility of applying RAPD markers to germplasm characterization and identification of U. pinnatifida gametophytes, and to provide a molecular basis for Undaria breeding.     

Spatially variable synergistic effects of disturbance and additional nutrients on kelp recruitment and recovery

Understanding the impact of multiple stressors on ecosystems is of pronounced importance, particularly when one or more of those stressors is anthropogenic. Here we investigated the role of physical disturbance and increased nutrients on reefs dominated by the canopy-forming kelp Ecklonia radiata. We combined experimental kelp canopy removals and additional nutrient at three different locations in a large embayment in temperate southeastern Australia. Over the following winter recruitment season, Ecklonia recruitment was unaffected by increased nutrients alone, but tripled at all sites where the canopy had been removed. At one site, the combination of disturbance and increased nutrients resulted in more than four times the recruitment of the introduced kelp Undaria pinnatifida. Six months after disturbance, the proliferation of the Undaria canopy in the canopy-removal and nutrient-addition treatment negatively influenced the recovery of the native kelp Ecklonia. Given the otherwise competitive dominance of adult Ecklonia, this provides a mechanism whereby Undaria could maintain open space for the following recruitment season. This interplay between disturbance, nutrients and the response of native and invasive species makes a compelling case for how a combination of factors can influence species dynamics.     

The structure of biogenic habitat and epibiotic assemblages associated with the global invasive kelp <Emphasis Type="Italic">Undaria pinnatifida</Emphasis> in comparison to native macroalgae

Kelp forests dominate temperate and polar rocky coastlines and represent critical marine habitats because they support elevated rates of primary and secondary production and high biodiversity. A major threat to the stability of these ecosystems is the proliferation of non-native species, such as the Japanese kelp Undaria pinnatifida (‘Wakame’), which has recently colonised natural habitats in the UK. We quantified the abundance and biomass of U. pinnatifida on a natural rocky reef habitat over 10 months to make comparisons with three native canopy-forming brown algae (Laminaria ochroleuca, Saccharina latissima, and Saccorhiza polyschides). We also examined the biogenic habitat structure provided by, and epibiotic assemblages associated with, U. pinnatifida in comparison to native macroalgae. Surveys conducted within the Plymouth Sound Special Area of Conservation indicated that U. pinnatifida is now a dominant and conspicuous member of kelp-dominated communities on natural substrata. Crucially, U. pinnatifida supported a structurally dissimilar and less diverse epibiotic assemblage than the native perennial kelp species. However, U. pinnatifida-associated assemblages were similar to those associated with Saccorhiza polyschides, which has a similar life history and growth strategy. Our results suggest that a shift towards U. pinnatifida dominated reefs could result in impoverished epibiotic assemblages and lower local biodiversity, although this could be offset, to some extent, by the climate-driven proliferation of L. ochroleuca at the poleward range edge, which provides complex biogenic habitat and harbours relatively high biodiversity. Clearly, greater understanding of the long-term dynamics and competitive interactions between these habitat-forming species is needed to accurately predict future biodiversity patterns.     

Undaria pinnatifida: A case study to highlight challenges in marine invasion ecology and management

Marine invasion ecology and management have progressed significantly over the last 30 years although many knowledge gaps and challenges remain. The kelp Undaria pinnatifida, or “Wakame,” has a global non‐native range and is considered one of the world's “worst” invasive species. Since its first recorded introduction in 1971, numerous studies have been conducted on its ecology, invasive characteristics, and impacts, yet a general consensus on the best approach to its management has not yet been reached. Here, we synthesize current understanding of this highly invasive species and adopt Undaria as a case study to highlight challenges in wider marine invasion ecology and management. Invasive species such as Undaria are likely to continue to spread and become conspicuous, prominent components of coastal marine communities. While in many cases, marine invasive species have detectable deleterious impacts on recipient communities, in many others their influence is often limited and location specific. Although not yet conclusive, Undaria may cause some ecological impact, but it does not appear to drive ecosystem change in most invaded regions. Targeted management actions have also had minimal success. Further research is needed before well‐considered, evidence‐based management decisions can be made. However, if Undaria was to become officially unmanaged in parts of its non‐native range, the presence of a highly productive, habitat former with commercial value and a broad ecological niche, could have significant economic and even environmental benefit. How science and policy reacts to the continued invasion of Undaria may influence how similar marine invasive species are handled in the future.     

Effects of the alien algae <Emphasis Type="Italic">Undaria pinnatifida</Emphasis> (Phaeophyceae,Laminariales) on the diversity and abundance of benthic macrofauna in Golfo Nuevo (Patagonia,Argentina): potential implications for local food webs

Changes in macroalgal community structure caused by invasive seaweeds have strong impacts on the associated macrofauna due to the role of macroalgae as autogenic ecosystem engineers. This study investigates the effect of Undaria pinnatifida on the abundance and diversity of benthic macrofauna in Golfo Nuevo, northern Patagonia, using a manipulative experiment involving the systematic removal of Undaria microthalli from strips of hard bottom during its eighth-month growing period. Species richness and diversity were higher in plots covered by Undaria than when Undaria was removed. Also, the abundance of two species of crustaceans, one species of sea urchin, one species of nemertina and several species of polychaetes was higher. We attribute these effects to the provision of new habitat structure by Undaria, a larger and structurally more complex species than the local native seaweeds. These results support the hypothesis that complex habitats enhance abundance and species richness, by offering different shelter and foraging opportunities compared to morphologically simpler habitats. Based on a review of diet studies in the region, we speculate that Undaria could potentially produce a bottom-up effect on local food chains by increasing abundance of prey for a wide variety of predators, from invertebrates to marine mammals. While our study has a narrow temporal and spatial scale, we expect similar effects of Undaria on the macrofauna in other sites of the Argentine coast, especially those dominated by small native macroalgae.     

Development and cultivation of F2 hybrid between Undariopsis peterseniana and Undaria pinnatifida for abalone feed and commercial mariculture in Korea

The kelp Undariopsis peterseniana (Kjellman) Miyabe et Okamura is warm water tolerant and consequently there is currently considerable interest in developing commercial cultivation techniques for this species in Korea. Undaria is an important species for both the abalone industry and for commercial seaweed mariculture. In an attempt to extend the culture period of Undaria we bred and cultured hybrid kelp that is a cross between free-living gametophytes of U. peterseniana and Undariopsis pinnatifida. The morphological characteristics of the F₁ hybrid sporophytes were intermediate between those of the parent plants having shallow pinnated blades and forming both sorus and sporophyll. A F₂ generation was produced by isolating zoospores from sorus and sporophyll separately from a F₁ hybrid thallus. Using free-living gametophyte seeding and standard on-growing techniques, F₀ (female U. pinnatifida and male U. peterseniana), F₁, and F₂ gametophytes were cultured from December 2011 to May 2013. The morphological differences between the F₁ and F₂ generations were assessed. The F₂ hybrids were found to have longer pinnate blades and narrower midribs than the F₁ hybrid and only formed sporophylls. Growth and morphology of F₂ hybrids originating from the sporophyll or sorus of the F₁ hybrids were not morphologically different from each other. Both of the F₂ hybrids exhibited late maturation, with the early stages of sporophylls appearing in April. This late maturation of F₂ hybrids is beneficial in the development of hybrid strains that extend the period of availability of Undaria for abalone feed and cultivation in Korea.     

BIOGEOGRAPHY AND ECOLOGY OF THE KELP/RED ALGAL SYMBIOSIS

A kelp/red algal symbiosis is described from nature based on extensive collections from the San Juan Islands, Washington. Kelp gametophytes were found as endophytes in the cell walls of seventeen species of red algae in three different kelp communities. Host red algae were mostly filamentous (e.g., Pleonosporium vancouverianum) or polysiphonous (e.g. Polysiphonia paniculata). The kelp gametophytes completed vegetative and reproductive development in the hosts with gametangia formed at the host surface and with sporophytes up to several mm in height being produced while still attached to the host. To date, none of the kelp gametophytes from nature have been identified to genus or species, although the gametophyte of Nereocystis luetkeana is a potential candidate for the symbiosis. Preliminary observations from Nova Scotia and the Isle of Man have not found the association in the Atlantic Ocean. Laboratory studies in Korea successfully reconstructed the symbiosis in the red alga Aglaothamnion oosumiense using zoospores of Undaria pinnatifida but not Laminaria religiosa. Here we outline the development of the symbiosis and discuss the potential adaptive significance of the kelp/red algal interaction.     

Algal communities at Gouqi Island in the Zhoushan archipelago, China

The kelp bed of Gouqi Island lies in the east of the East China Sea, in the Shengsi Archipelago. It is a key component of the island-reef ecosystem. This study evaluated kelp beds on intertidal and subtidal zones from 2004 to 2006. We evaluated seasonal variations in the community structure of macroalgae, and their relationship with water temperature, nitrogen and phytoplankton. The Gouqi Island kelp bed consists mainly of species of Sargassum. Sargassum horneri was the dominant species in subtidal zones, representing 90% of the total biomass. The dominant species in intertidal zones were S. fusiforme, S. horneri, S. thunbergii and Undaria pinnatifida. Fifty-four phytoplankton species were found in the Gouqi Island kelp bed, of which 51 species were found inside the kelp beds, 43 species outside the kelp beds, and 40 species were present both inside and outside the kelp beds. Diatoms were dominant both inside and outside the kelp beds. Except in winter, Skeletonema costatum was dominant, and its abundance in autumn was over 98% of the total abundance. Water temperature was found to be the primary factor influencing the growth of S. horneri in the Gouqi Island kelp bed. S. horneri grew slowly from November to March, and rapidly from March to June. The upper temperature limit for growth of S. horneri was about 18°C. Higher water temperatures can result in a decline in the kelp bed. Nitrogen seemed to be a limiting factor for macroalgal growth. This was particularly true for S. horneri and phytoplankton. Since nitrate is the primary nutrient for S. horneri, its absorption by S. horneri resulted in seasonal changes of nitrate in the ecosystem.     

Cultivation and utilization of Undaria pinnatifida (wakame) as food

A brief review is presented concerning wakame, Undaria pinnatifida, one of the most popular seaweeds used for food in Japan. Although it has been cultivated since about 1940, full-scale cultivation occurred after 1955. As methods for providing ‘seed stock’ and of processing the harvested sporophytes progressed, the yield increased rapidly. The main areas of cultivation are in Japan (e.g. Sanriku, Naruto), Korea and China, while ‘wild’ U. pinnatifida has been introduced into France, New Zealand and Australia. The total world yield of wakame exceeds 500 000 t fresh weight. Cultivated and harvested Undaria is boiled and salted (thus becoming green) and refrigerated; in the factory, it is removed its foreign matter and salt and dried. After checking for quality, the product is packaged in forms convenient for cooking and eating.