Sargassum muticum

Invasion History

First Non-native North American Tidal Record: 1944
First Non-native West Coast Tidal Record: 1944
First Non-native East/Gulf Coast Tidal Record:

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General Invasion History:

Sargassum muticum was first described from Wakayama Prefecture, Japan, by Yendo in 1907. Its native range extends from the East China Sea (China and South Korea) to the coasts of Japan and the north shore of Hokkaido, and the Sakhalin and Kurile Islands, Russia (Critichley 1983b; Eneglen et al. 2015). It is introduced in the Eastern Pacific and Eastern Atlantic and genetic studies indicate that these populations probably originated from the Seto Inland Sea, Japan (Cheang et al. 2010). In North America, S. muticum was first introduced to Vancouver Island, British Columbia, where it was discovered in 1944, probably introduced with Pacific Oysters (Crassostrea gigas) from Japan. By 1977, it had spread north to Ketchikan, Alaska (Scagel et al. 1956; Scagel et al. 1989). The pattern of spread/discovery to the south was rather jumpy, with first records in Coos Bay, Oregon in 1947; Mission Bay, California (CA) in 1959; San Francisco Bay, CA in 1963; and Los Angeles, CA in 1973. By the 2000s it had colonized most of the Pacific coast of Baja California (Scagel et al. 1956; Scagel et al. 1989; Engelen et al. 2015). In Europe, it was first collected in Southeast England in 1971 and rapidly spread north and south, reaching Norway by 1988, Spain by 1985, and Morocco by 2012 (Farnham 1980; Runess 1989; Fernandez et al. 1990; Sabour et al. 2013). In 1980, it was first collected in lagoons on the Mediterranean coast of France and by 1992 it was established in the Lagoon of Venice (Knoepfller-Peguy et al. 1985; Curiel et al. 1998), but its distribution in the Mediterranean Sea remains localized (Engelen et al. 2015; Thibaut et al. 2015). The large size of this seaweed, its rapid spread, and high abundances in many locations, have led to the extensive study of its ecology and impacts (Engelen et al. 2015).

North American Invasion History:

Invasion History on the West Coast:

Sargassum muticum was first collected on the West Coast of North America, at White Rock and Buccaneer Bay, north of Vancouver, British Columbia in 1944, in sites where Pacific Oysters where cultivated. It was initially mistaken for the native seaweed Cytoseira geminata. By 1952, it was found at many sites in the Strait of Georgia and Puget Sound, including the San Juan Islands and the Strait of Juan de Fuca (Scagel 1956). The pattern of discovery along the coast of Oregon and Washington suggests several independent introductions with oyster transplants, with subsequent spread by drifting weed and fouled boats. It was found in Willapa Bay, Washington in 1953; Oceanside Beach (near Tillamook Bay), Oregon, in 1951; and Coos Bay, OR in 1947 (Scagel 1956). Spread to the north was slower, but it was found near Ketchikan, Alaska in 1974-1977 (Scagel et al. 1989; Engelen et al. 2015; University of Alaska Southeast Herbarium 2016), and Haida Gwaii, British Columbia in 1981 (Sloan and Bartier 2004).

The first record of S. muticum in California is a report of its occurrence in Mission Bay, San Diego in 1958, which looks like a sudden jump (Stewart 1991), possibly by ship transport. Other records are consistent with a progressive march to the south: Crescent City Harbor in 1963 (Norton 1981, Cohen 2005); Humboldt Bay in 1965 (Dawson 1965, cited by Boyd et al. 2002); Tomales Bay and San Francisco Bay in 1973 (Cohen and Carlton 1995); Monterey Bay in 1977 (Carlton et al. 1996; Cohen 2005); Santa Barbara in 1977 (Cohen 2005); Los Angeles in 1973; (Norton 1981); and San Diego Bay in 1969 (Cohen 2005). The rapid spread is surprising, since S. muticum requires sheltered waters for establishment (Norton 1981). By 1973, it was already established in Ensenada, Mexico (Norton 1981) and by 1984 it had reached Punta Abreojos, Baja California, Mexico (Espinosa 1990). Overall, in 70 years since its invasion on the West coast, it has colonized ~30 degrees of latitude and a span of 4000 km of coastline (Engelen et al. 2015).

Invasion History Elsewhere in the World:

In the Northeast Atlantic, S. muticum was first found on Portsea Island, Portsmouth, England in 1971, and rapidly spread through the Solent, between the south coast of England and the Isle of Wight (Farnham 1980). It colonized the French side of the English Channel by 1977 (Gruet 1977, cited by Farnham 1980). This seaweed reached the Netherlands by 1980, the German portion of the Wadden Sea by 1983 (Buschbaum et al. 2012), and the western mouth of the Limfjord, Denmark, by 1984 (Staer et al. 2000). Sargassum muticum colonized the Kattegat in Denmark and Sweden, the westernmost portion of the Baltic Sea, in 1993-1996, probably reaching the lower limit of its salinity tolerance (Karlsson and Loo 1999; Thomsen et al. 2007). Moving north, S. muticum spread into Norwegian and Swedish waters of the Skaggerak in 1987-1988 (Rueness 1989; Karsson and Loo 1999) and north up the west coast of Norway, to Bergen in 1989 (Hopkins 202) and Sogn og Fjordane (Bjærke and Fredriksen 2005). To the south, this seaweed reached Roscoff, at the tip of Brittany in 1980 (Belsher and Pommellec 1988), and reached the Basque Province of Spain by 1985 (Fernandez et al. 1990), and Portugal by 1989 (Chainho et al. 2015). In 2012, S. muticum was collected at several sites on the Atlantic coast of Morocco. The southernmost was Oualidia at 32°N (Sabour et al. 2013). In ~40 years, S. muticum has spread over 39 degrees of latitude and a 5600 km span of coastline (Engelen et al. 2015).

In contrast with its spread on the West Coasts of Europe and North America, Sargassum muticum's spread in the Mediterranean has been limited, spotty, and punctuated by extinctions. In 1985-1990, S. muticum was found in 14 lagoons on the French Mediterranean Coast, but in surveys in 2012 established populations were found in only four lagoons (Thibaut et al. 2015). In 1992, established populations of the seaweed were found in the Venice Lagoon. Dense populations occur on wharves and breakwaters (Curiel et al. 1998). Sargassum muticum was found in a lagoon on the east coast of Corsica, but was not seen in later surveys (Thibaut et al. 2015). Some floating specimens of this seaweed have been found in the open Mediterranean off Spain, but S. muticum has only become established in lagoons with limited exchange (Engelen et al. 2015).

Culture of the Pacific Oyster (Crassostrea gigas) has been the initial vector for the three major invasions of S. muticum in the Northeast Pacific and Atlantic, and the Mediterranean (Scagel 1956; Farnham 1980; Knoepffler-Peguy et al. 1985). Regional and local dispersal could occur by hull fouling of commercial ships or recreational boats and by the natural dispersal of floating mats of seaweed (Engelen et al. 2015).

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Description

Sargassum muticum usually grows upright in the water column, attached by a solid, conical, spongy holdfast about 50 mm across. The holdfast gives rise to a tapered axis up to 50 mm in height. In young plants, leaves, up to 200 mm long, arise from the main axis. These basal leaves have a well-defined mid-rib. In older plants, the main axis may branch, making the plant bushier. Lateral shoots grow off the main axis in spirals with scale-like triangular leaves at the base, which protect the growing bud. The main axis is perennial, while the lateral shoots die off after the growing season. Primary lateral stems are cord-like, usually growing to 1-3 m, but sometimes 6-10 m. In populations on the West Coast of Ireland, mature plants had 40-100 primary lateral branches (Baer and Stengel 2010), while a population in the Netherlands had 1-16 (Critchley et al. 1987). Secondary lateral stems arise off the primary laterals, branching off in the axil of a leaf. As the lateral stems grow, they become twisted due to water movement. Leaves from the primary and secondary laterals, produced in the winter, are small, less than 200 mm long, thick, and have a weak mid-rib. During the summer, the fronds become fertile and the new leaves are thinner, narrow, and lacking a midrib. They have an uneven outline, resembling holly leaves. The lateral branches bear air-bladders and reproductive receptacles in the axils of the leaves. The air-bladders are spherical to pear-shaped, on stalks, about 3 mm in diameter. The receptacles vary in shape from elliptical to cylindrical to spindle-shaped and are usually 20--30 mm long (sometimes up to 60 mm). The plants are yellow-green to olive-brown. This description is based on: Abbott and Hollenberg 1976; Critchley 1983a; Cohen 2005; and Engelen et al. 2015.

Sargassum muticum is one of several similar species known in Japan, and its taxonomic history is complex. It was initially described as a variety of S. kjellmanianum, which is now regarded as a synonym of S. miyabe (Critchley 1983b; Cheang et al. 2010; Engelen et al. 2015). The genetic diversity of introduced populations in North America and Europe is low and all populations are close to S. muticum populations from western and central Japan (Cheang et al. 2010).

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Taxonomy

Ecology

General:

Sargassum muticum is monoecious and mature plants produce receptacles, which contain both male and female conceptacles for the production of sperm and eggs (Bold and Wynne 1978; Engelen et al. 2015). Receptacles can constitute 24-55% of the plant's biomass, but this reproductive output is fairly typical for fucoid seaweeds (Engelen et al. 2015). Gamete release is synchronized in a semilunar, 14-day cycle, with release peaking at the full and new moons, just after spring tides (Engelen et al. 2008; Engelen et al. 2015), though with some variation with local cues (Monteiro et al. 2009b). Eggs are fertilized in the receptacles, but are retained, attached to the receptacle surface for a few days and are then released as germlings that sink quickly. The propagules usually settle within a few meters of the parents, but can be found up to 1.3 km away. They could potentially disperse over longer distances, since they retain the ability to settle for 49 days (Deysher and Norton 1982). Each receptacle releases ~ 300 propagules and a small plant can release up to 500,000 propagules (Norton and Deysher 1989, cited by Engelen et al. 2015). The fertilized zygotes average about 0.25 mm in diameter (Deysher and Norton 1982). Germlings have comparatively large, basal leaves, with developing lateral axes (Critchley 1983a). Breeding seasons vary geographically, peaking in spring-summer in Northern Europe and Northwestern North America, but with a longer season at southern locations such as Portugal, southern California, and Mexico (Deysher 1984; Aguilar-Rosas and Galindo 1990; Engelen et al. 2015). After releasing their propagules, adult plants lose their fronds and disintegrate, except for the perennial holdfast, which gives rise to new fronds in the next growing season (Critchley 1983a; Engelen et al. 2015). This dormant period is usually in autumn in the northern part of the range, but occurs in summer-fall in southern Portugal, California, and the Venice Lagoon (Deysher 1984; Sfriso and Facca 2013; Engelen et al. 2015).

Sargassum muticum grows over a wide latitudinal range, from cold-temperate to subtropical conditions, from temperatures below 0°C in Sweden (Karlsson and Loo 1999) to 30°C in the Venice Lagoon (Sfriso and Facca 2013). In the laboratory, germlings survived at salinities as low as 6.8 PSU, but with minimal growth (Hales and Fletcher 1989; Karlson and Loo 1999; Steen 2004; Thomsen et al. 2007). Germlings are more sensitive to temperature and salinity than adult plants and grow successfully between 15 and 25°C and 15-35 PSU (Hales and Fletcher 1989; Steen 2004). Early germlings (2 weeks old) showed increasing growth at 9 to 44 µE m-3s-1 and had decreased growth at 88 µE m-3s-1, while older germlings grew well at 18 to 88 µE m-3s-1, and were more tolerant of high light (Hales and Fletcher 1989). Sargassum muticum is not a good competitor at low light and has a strategy of growing fast to form a canopy in shallow water. It can adjust its pattern of growth and branching according to the density of its neighbors, permitting it to form very dense populations (Engelen et al. 2015).

In its native range in Asia, S. muticum occurs on rocky shores from the lower intertidal to 4 m depth and is not known as an aggressive colonizer of artificial structures (Engelen et al. 2015). It is generally associated with sheltered locations. Plants transplanted to exposed locations suffered increased breakage and lower growth rates (Viejo et al. 1995). In invaded habitats, it occurs on pilings, floats, marinas, in canals, aquaculture cages, marinas, breakwaters, oyster beds, and eelgrass beds (Belsher and Pommellec 1988; den Hartog 1997; Harries 2007; Kraan 2008; Engelen et al. 2015). In Strangford Lough, Northern Ireland, S. muticum has colonized soft-sediment habitats by 'stone-walking' on rock fragments and shells and spreading by water movements (Strong et al. 2006). Adult plants, dislodged by waves or disturbance, or fragments of degenerating plants with fertile receptacles, form floating mats (Norton 1981). Sargassum muticum, like other fucoid seaweeds, produces organic compounds for chemical defense against microbes, epiphytes, and grazers. Phenolic compounds are greatest during the reproductive period, possibly for protection of the receptacles (Plouguerne et al. 2006; Plougerne et al. 2008). In spite of these compounds, S. muticum supports a dense community of epiphytes and epifauna, and is grazed by a variety of invertebrates, with varying degrees of preference (Norton and Benson 1983; Monteiro et al. 2009; Strong et al. 2009; Baer & Stengel 2010; Cacabelos et al. 2010; Gestoso et al. 2010).

Consumers:

snails, amphipods, sea urchins

Competitors:

Native seaweeds, seagrasses

Trophic Status:

Primary Producer

PrimProd

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Habitats

General Habitat	Marinas & Docks	None
General Habitat	Rocky	None
General Habitat	Oyster Reef	None
General Habitat	Grass Bed	None
General Habitat	Canals	None
General Habitat	Unstructured Bottom	None
Salinity Range	Mesohaline	5-18 PSU
Salinity Range	Polyhaline	18-30 PSU
Salinity Range	Euhaline	30-40 PSU
Tidal Range	Subtidal	None
Tidal Range	Low Intertidal	None
Vertical Habitat	Epibenthic	None
Vertical Habitat	Littoral	None

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Life History

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	0	Field observations, Sweden (Karlsson and Loo 1999)
Maximum Temperature (ºC)	30	Highest tested, lab experiments (Hales and Fletcher 1989)
Minimum Salinity (‰)	10	Lab experiments- Growth greatly reduced compared to 27-34 PSU (Hales and Fletcher 1989)
Maximum Salinity (‰)	35	Highest tested, lab experiments (Hales and Fletcher 1989)
Minimum Reproductive Temperature	15	Experimental (Kerrison and Le 2016)
Maximum Reproductive Temperature	25	Experimental (Kerrison and Le 2016)
Minimum Reproductive Salinity	20	Minimum for fertilization. Germlings can tolerate brief exposure to 5 PSU, especially in 2nd and 3rd weeks after fertilization (Steen 1984)
Maximum Reproductive Salinity	40	Experimental, germlings in receptacle (Kerrison and Le2016)
Minimum Length (mm)	300	Moyrus, Ireland, stunted plants at exposed site, but 100% fertile (Baer and Stengel 2010)
Maximum Length (mm)	10,000	Engelen et al. 2015
Broad Temperature Range	None	Cold temperate-Warm temperate
Broad Salinity Range	None	Mesohaline-Euhaline

General Impacts

Sargassum muticum is one of many Sargassum species in its native Asian range, and is not especially prominent or aggressive there. In the Northeast Pacific and Northeast Atlantic, however, it has spread rapidly along coastlines and has become a major component of algal communities (Engelen et al. 2015). Economic impacts to beach use, boating, power plants, fisheries, and aquaculture, were reported in the early years of the invasion in England and France. These impacts led to a number of unsuccessful eradication attempts (Critchley et al. 1986; Belsher and Pommellec 1989). Ecological impacts of S. muticum have been extensively studied, particularly along the coasts of northern Europe, northern Spain and Portugal, and the San Juan Islands, in northern Puget Sound, Washington. These include competition for space, light, and nutrients, alterations of habitat and associated epibiota, and effects on grazers (Schaffelke and Hewitt 2007; Engelen et al. 2015).

Economic Impacts

Early accounts of the spread of Sargassum muticum on the West Coast of North America (Scagel 1956; Norton 1981) make no mention of adverse economic impacts. A later survey, focused on Sargassum horneri and Undaria pinnatifida refers to S.muticum as 'naturalized' and does not refer to economic impacts (Kaplanis et al. 2016). In Southeastern England and Atlantic France, extensive impacts, including fouling of boats, fishing nets, aquaculture equipment, and power plant intakes, was reported, together with large floating mats in nearshore waters and rotting masses piling up on beaches (Critchley et al. 1986; Belsher and Pommellec 1988). In 1974-1975, an eradication was attempted in England with large volunteer groups, hand-picking intertidal plants. The volunteers collected 31 metric tons, but missed germlings and juvenile plants and eradication was found to be impractical. Trawls, cutting machines, and suction machines were tested as alternatives to hand-picking, but considered un-economical. Sargassum muticum was considered unsuitable for industrial and food products (Critchley et al. 1986), although it is eaten in Korea. A recent review of potential economic uses found that use of 'wild' plants in Europe for biofuel, food, or fertilizer was unsafe because of the accumulation of heavy metals and high ash content. However, S. muticum contains fucoxanthins, antioxidants, and other compounds of pharmaceutical interest (Milledge et al. 2106).

Ecological Impacts

Competition- Interactions between S. muticum and native biota, in many different marine communities have been studied, including rocky shores, tidepools, kelp beds, seagrass meadows, and soft-bottom communities (Schaffelke and Hewitt 2007; Engelen et al. 2008). Competitive interactions have been especially noted with the algae of the closely related perennial genera Cytoseira and Halidrys spp. Sargassum’s semi-perennial life-style, dying back to the holdfast, in unfavorable seasons may be a competitive advantage. In Portugal, its dying fronds denied space to C. humlis (Engelen and Santos 2009). In Denmark, S. muticum outgrew Halidrys siliqua, possibly because it did not need to invest in structural strength for winter survival (Wernberg et al. 2000). Sargassum muticum's bushy growth form and buoyant branches permit it to form a canopy and shut out light and occupy space during the most favorable time for growth (Curiel et al. 1998; Britton-Simmons 2004; Harries et al. 2007; Salvaterra et al. 2013). However, S. muticum often seems to require disturbance to invade established communities (de Wreede 1983, cited by Schaffelke and Hewitt 2007; Strong and Dring 2011; Bertocci et al. 2014). Sargassum muticum invaded rocky areas off California and in the San Juan Islands after die-offs or experimental removal of kelps (Ambrose and Nelson 1982; Britton-Simmons and Abbott 2008).

Habitat Change- When S. muticum replaces native seaweeds and seagrasses, it can alter the structure of native habitats. In many cases, S. muticum supports a similar or a more abundant and diverse epiphytic and epifaunal community than the native flora (Viejo 1999; Buschbaum et al. 2006; Engelen et al. 2015). Its ability to colonize bare, soft substrate by 'stone-walking' on shells and stones, and to colonize Eelgrass (Zostera marina) beds greatly increases the structural complexity of these habitats (den Hartog 1997; Strong et al. 2011; DeAmicis et al. 2015). It is used as habitat by fishes, crustaceans, and cuttlefish (Engelen et al. 2015; Stiger-Pouvreau and Thouzeau 2015).

Food/Prey- Herbivores vary considerably in their response to S. muticum. Strong avoidance was noted for sea urchins (Strongylocentrotus droebachiensis) in northern Puget Sound (Britton-Simmons 2004) and Psammechinus miliaris in Denmark (Pedersen et al. 2005). Other grazers showed lesser levels of avoidance, or no preference (Monteiro et al. 2009; Cacabelos et al. 2010; Engelen et al. 2011). Some grazers appear to feed heavily on S. muticum, such as the amphipod Dexamine spinosa in Strangford Lough, Northern Ireland (Strong et al. 2009), and the snail Lacuna vincta in Northern Puget Sound. The latter seems to have developed its preference in the last 30 years (Britton-Simmons et al. 2011). In many of the systems where it has been studied, S. muticum seems to be under light grazing pressure (Norton and Benson 1983; Pedersen et al. 2005; Monteiro et al. 2009; Cacabelos et al. 2010; Engelen et al. 2011; Mabey et al. 2022). However, because of its rapid growth, large biomass, and seasonal die-offs, it is a major contributor to primary production in many intertidal and subtidal systems, and to subtidal detritus and intertidal wrack (Rossi et al. 2009; Olabarria et al. 2010; Vaz-Pinto et al. 2014; Engelen et al. 2015).

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Regional Impacts

NEP-VI	Pt. Conception to Southern Baja California		Ecological Impact	Competition
Sargassum muticum colonized Giant Kelp (Macrocystis pyrifera) beds on Bird Rock, off Catalina Island, after a die-off, possibly caused by high water temperatures during El Nino. Shading by Sargassum muticum probably inhibited recolonization by the kelp. After experimental removal, the kelp recolonized the cleared areas (Ambrose and Nelson 1982). Several years later, Giant Kelp did recolonize Bird Rock (Foster & Schiel 1992, cited by Engelen et al. 2015). Removal experiments in tidepools at Little Corona del Mar in Newport Beach in southern California, showed little effect level, pool temperature, seaweed biomass and community composition, or faunal composition. Recovery of S. muticum populations was rapid (Smith 2016).
P058	_CDA_P058 (San Pedro Channel Islands)		Ecological Impact	Competition
Sargassum muticum colonized Giant Kelp (Macrocystis pyrifera) beds on Bird Rock, off Catalina Island, after a die-off, possibly caused by high water temperatures during El Nino. Shading by Sargassum muticum probably inhibited recolonization by the kelp. After experimental removal, the kelp recolonized the cleared areas (Ambrose and Nelson 1982). Several years later, Giant Kelp did recolonize Bird Rock (Foster & Schiel 1992, cited by Engelen et al. 2015).
P040	Newport Bay		Ecological Impact	Competition
Removal experiments in tidepools at Little Corona del Mar in Newport Beach in southern California, showed little effect level, pool temperature, seaweed biomass and community composition, or faunal composition. Recovery of S. muticum populations was rapid (Smith 2016).
P090	San Francisco Bay		Ecological Impact	Habitat Change
In field cage experiments, juvenile Chinook Salmon (Oncorhynchus tshawystcha) were reared in several habitats, bare sand, Eelgrass (Zostera marina, Sargassum muticum and mixed Eelgrass-Sargassum. Growth was best in the mixed habitat, suggesting that habitat variety is important (Hughes et al. 2020).
NEP-V	Northern California to Mid Channel Islands		Ecological Impact	Habitat Change
In field cage experiments, juvenile Chinook Salmon (Oncorhynchus tshawystcha) were reared in several habitats, bare sand, Eelgrass (Zostera marina, Sargassum muticum and mixed Eelgrass-Sargassum. Growth was best in the mixed habitat, suggesting that habitat variety is important (Hughes et al. 2020).
CA	California		Ecological Impact	Competition
Sargassum muticum colonized Giant Kelp (Macrocystis pyrifera) beds on Bird Rock, off Catalina Island, after a die-off, possibly caused by high water temperatures during El Nino. Shading by Sargassum muticum probably inhibited recolonization by the kelp. After experimental removal, the kelp recolonized the cleared areas (Ambrose and Nelson 1982). Several years later, Giant Kelp did recolonize Bird Rock (Foster & Schiel 1992, cited by Engelen et al. 2015)., Removal experiments in tidepools at Little Corona del Mar in Newport Beach in southern California, showed little effect level, pool temperature, seaweed biomass and community composition, or faunal composition. Recovery of S. muticum populations was rapid (Smith 2016).
CA	California		Ecological Impact	Habitat Change
In field cage experiments, juvenile Chinook Salmon (Oncorhynchus tshawystcha) were reared in several habitats, bare sand, Eelgrass (Zostera marina, Sargassum muticum and mixed Eelgrass-Sargassum. Growth was best in the mixed habitat, suggesting that habitat variety is important (Hughes et al. 2020)., In field cage experiments, juvenile Chinook Salmon (Oncorhynchus tshawystcha) were reared in several habitats, bare sand, Eelgrass (Zostera marina, Sargassum muticum and mixed Eelgrass-Sargassum. Growth was best in the mixed habitat, suggesting that habitat variety is important (Hughes et al. 2020).

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