Grateloupia turuturu

Invasion History

First Non-native North American Tidal Record: 1996
First Non-native West Coast Tidal Record: 2008
First Non-native East/Gulf Coast Tidal Record: 1996

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General Invasion History:

Grateloupia turuturu is native to the Northwest Pacific from Vladivostok, Russia to Jeju Island, South Korea (Gavio and Frederiq 2002; Lee et al. 2009; Yang and Kim 2015; Guiry and Guiry 2015). In 1969, it was noticed in the Solent Estuary, England and initially identified as G. doryphora, described from Peru (Farnham 1980). This alga spread throughout European waters, reaching Brittany in 1988 (Goulletquer et al. 2002); the Netherlands in 1993 (Marston and Villalard-Bohnsack 2002; Stegenga and Karremans 2015); Galicia, Spain in 1991 (Barbara and Cremades 2004); and northern Portugal in 1997 (Araújo et al. 2003, cited by Barbara and Cremades 2004). In the Mediterranean it was first found in the Thau Lagoon, France (Verlaque 2001), and now occurs at sites from the Alboran Sea, Spain (reported in 1988, Barbara and Cremades 2004), to the Venice Lagoon (reported in 1992, Tolomio 1993) and Israel (reported in 2013, Katsanevakis et al. 2014). Gavio and Fredericq (2002) used genetic techniques to demonstrate that this alga was, in fact, G. turuturu.

Grateloupia turuturu was first reported (as G. doryphora) from Narragansett Bay, Rhode Island in 1996, and is now known from Boston Harbor to Long Island (Villalard-Bohnsack and Harlin 1997; Van Patten 2006; Mathieson et al. 2007). On the West Coast, it is established in Pillar Point Harbor, outside San Francisco Bay, and in Huntington Harbor, near Los Angeles (Zabin et al. 2011; California Department of Fish and Wildlife 2014). Other widely scattered and genetically confirmed invasions are in the Azores (reported in 2007, Micael et al. 2014; Chainho et al. 2015); Madeira (reported in 2002, Ferreira et al. 2012, cited by Chainho et al. 2015); Brazil (reported in 2011, Araujo de Azevedo et al. 2015); and New Zealand (reported in 2005, D'Archino et al. 2007, 2 specimens). Recognition of these invasions is complicated by occurrence of native species of Grateloupia in different regions. Reports from several locations on the west coast of Africa from the Canary Islands to Namibia (John et al. 2004, cited by Guiry and Guiry 2016) need to be checked by genetic methods.

North American Invasion History:

Invasion History on the West Coast:

An established population of Grateloupia turuturu was first discovered on the West Coast of North America in Ensenada, Baja California, Mexico in 2008 (Miller et al. 2011; Aguilar-Rosas et al. 2012). In 2009, specimens of G. turuturu were found attached to docks in the Santa Barbara yacht harbor (Hughey et al. 2009). This alga was collected in 2010 in Pillar Point Harbor, Half Moon Bay (Miller et al. 2011; Zabin et al. 2011). In 2011, G. turuturu was found in two locations, Huntington Harbor and Mission Bay (California Department of Fish and Wildlife 2014). The identification of specimens in Mexico and Santa Barbara was confirmed by molecular comparison with specimens from Japan (Hughey et al. 2009; Aguilar-Rosas et al. 2012). Most of these locations were in small yacht harbors. It is likely that these scattered populations stem from an initial ballast water or hull fouling introduction by ships to a commercial port (Hughey et al. 2009) with secondary spread by recreational boats to smaller bays and harbors.

Invasion History on the East Coast:

On the East Coast, Grateloupia turuturu was first collected in 1996 near Newport, Rhode Island, in Narragansett Bay, and initially identified as G. doryphora. In Narragansett Bay, it appears to be confined to the lower and middle portions of the estuary (Villalard-Bohnsack and Harlin 1997; Harlin and Villalard-Bohnsack 1999; MacIntyre et al. 2011). In the 20 years since its initial discovery, its spread has been relatively limited. In 2001, it was collected at Montauk, New York, at the tip of Long Island Sound (Gavio and Fredericq 2002), and in 2004 it was found at Millstone Point, Waterford, Connecticut and is established in the Sound (Van Patten 2006; Janiak and Whitlatch 2012). In 2007, it was collected in Buzzards Bay, Cape Cod Bay, and Boston Harbor, Massachusetts (MIT Sea Grant 2007; Mathieson et al. 2007). In 2014, it was found in Great Bay New Hampshire, and in 2017 it was found in the Damariscotta River estuary in Walpole, Maine (Capistrant-Fossa, and Brawley 2019). Ballast water or hull fouling are likely vectors for introduction; the first collections on the East Passage of Narragansett Bay were in a region of commercial ship traffic (Villalard-Bohnsack and Harlin 1997).

Invasion History Elsewhere in the World:

Grateloupia turuturu was first reported from Europe (as G. doryphora) from the Solent Estuary, off the English Channel. As of 1975, its range was still relatively limited to a small area of the southern coast of England, in the vicinity of the Isle of Wight (Farnham 1980). By the 1990s, it had spread west to the Fleet Lagoon, Dorset, and east to Jersey in the Channel Islands (Farnham 1994; Eno et al. 1997). On the European mainland, it was found in Lorient, on the south coast of Brittany, in 1988 (Goulletquer et al. 2002), and in Stavenisse, Netherlands in 1993 (Stegenga and Karremans 2015). In 1991, it was found in the Ria de Arousa, Galicia, on the northern Atlantic Coast of Spain (Barbara and Cremades 2004) and in northern Portugal (Chainho et al. 2015). The first record of G. turuturu from the Mediterranean was from the Thau Lagoon, France, a hotspot of oyster culture and seaweed introductions (Verlaque 2001). This seaweed has been spread to widely scattered locations in the Mediterranean, including Malaga, Spain in 1988 (Barbara and Cremades 2004); the Venice Lagoon in 1992 (Tolomio 1993); Mar Piccolo and Mar Grande of Taranto, Italy in 2001 (Gravili et al. 2010), and Zikim Beach, Israel in 2013 (Katsanevakis et al. 2014). In Europe, transplants of Pacific Oysters (Crassostrea gigas) have been a frequent vector of introduction, in addition to ballast water and hull fouling.

Grateloupia turuturu has been reported from the Azores and Madeira. Micael et al. (2014) and Chainho et al. (2015) give 1914 as the date of introduction for the Azores, probably as G. doryphora, but this may reflect taxonomic confusion. It was found in Madeira in 2002 (Ferreira et al. 2012). John et al. (2004, cited by Guiry and Guiry 2004) reported G. turuturu from the Canary Islands, Senegal, the Gambia, Liberia, Ghana, Angola, and Namibia. Since this alga's distribution is mostly temperate, these records need molecular confirmation. However, molecular data do confirm the identity of specimens collected in Florianopolis, Santa Catarina, Brazil in 2011 (de Azevedo et al. 2015).

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Description

Grateloupia turuturu is one of the largest known red algae. It grows as a large, flat, thick blade, or as clumps of up to eight blades, growing from a small (~5 mm) disk-shaped holdfast and a short, narrow, cylindrical stipe (stem). The blades vary in size and shape, from ribbon-like to kidney-shaped, and can be irregularly divided. The margins are wavy and often have thread-like or lance-shaped projections. The blades have a slippery and gelatinous texture. The blades are 140–600 µm thick and have an outer cortex of rounded or ovoid pigmented cells, and inner medulla of clear filaments formed from elongated cells. Size and shape of the blades vary greatly from 150–250 mm for kidney-shaped blades to 750–1000 mm for a more linear blade. Occasionally, plants have reached 2950 mm (2.95 m) in size (Simon et al. 2001, Brittany). New plants bud out asexually from the blade margins. The two major phases in the life cycle, diploid tetrasporophytes and gametophytes are morphologically similar. The tetrasporophytes have tetrasporangia scattered across the leaf. These are large ovoid cells which divide in a four-fold, cross-shaped (cruciate) pattern. In female gametophytes, the blades have numerous bottle-shaped ampullae, in which the carposporangium, which produces the carpospore or egg, is located. The color of the blade is deep red, burgundy, or maroon. Dying blades are yellowish. This description is based on: Villalard-Bohnsack and Harlin 1997, Harlin and Villalard-Bohnsack 1999, Simon et al. 2001, Gavio and Fredericq 2002, Van Patten 2006, and Mathieson et al. 2007.

Introduced forms of Grateloupia in British, French, and New England waters were initially identified as G. doryphora (Montagne) Howe, originally described from Peru (Farnham 1980; Tolomio 1993; Villalard-Bohnsack and Harlin 1997). Using molecular techniques, Gavio and Fredericq (2002) matched specimens from these locations with G. turuturu Yamada, native to the Northwest Pacific.

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Taxonomy

Ecology

General:

Grateloupia turuturu has a complex life cycle with two morphologically similar phases, a diploid tetrasporophyte and haploid gametophyte. A third stage, the carposporophyte, developed only within a specialized branch of the female thallus. The blades of the tetrasporophytes are covered with asexual tetrosporangia. The tetrasporangia divide in a fourfold (cruciate) pattern to produce tetraspores, ~20 µm in diameter. The tetraspores are released, and settle and grow into gametophytes. The blades of female thalli develop numerous carpogonia between the medulla and the cortex. The carposporophyte develops within an ampulla (bulb), formed by a system of filaments between the cortex and medulla of the thallus (Bold and Wynne 1978; Villalard-Bohnsack and Harlin 1997; Araujo et al. 2011). The carposporophyte produces a carpospore, the equivalent of an egg, which is fertilized and then released through the mouth of the bulb. In studies of two introduced populations, in Portugal (Araujo et al. 2011) and Rhode Island (Villalard-Bohnsack and Harlin 1997), male gametophytes were not found, so the role of sexual reproduction is unclear. In Rhode Island and Portugal, reproductive tetrasporophytes and gametophytes were found year-round (Villalard-Bohnsack and Harlin 1997; Araujo et al. 2011). Spores settle and grow as crusts. Filaments, initially one cell thick, arise from the crust and grow into upright blades. In lower Narragansett Bay, Rhode Island, G. turuturu reached 50-60% cover on boulders and bedrock, but dropped to low levels in early spring, before rapid growth and recruitment in May and June (Harlin and Villalard-Bohnsack 1997).

Grateloupia turuturu occurs in a wide range of coastal habitats. Most populations are known from cold-temperate to warm-temperate regions (Guiry and Guiry 2016), but it has been genetically confirmed in Brazil (Araujo de Azevedo et al. 2015) and reported from tropical Africa (John et al. 2004, cited by Guiry and Guiry 2004). The tropical records need to be confirmed by genetic means. In Brittany, G. turuturu was found over a salinity range of 15 to 38 PSU (Simon et al. 2001). It grows on a range of substrates, including bedrock, boulders, cobbles, shells, and artificial substrates, including ship hulls (Villalard-Bohnsack and Harlin 1997; Simon et al. 2001). This seaweed is found in the shallow subtidal, lower intertidal rocks, and tidepools (Villalard-Bohnsack and Harlin 1997; Simon et al. 2001). The crusts of G. turuturu grow over a wide range of irradiance (10-90 µE m-3s-1), but are more strongly affected by temperature (15-25°C), with no development at 10°C and development at the lowest light (10 µE m-3s-1) only at 25°C (Wei et al. 2013). In Brittany, G. turuturu was frequently associated with eutrophic conditions, including a sewage outfall (Simon et al. 2001). Older blades of Grateloupia turuturu were colonized with the red alga Polysiphonia fibrillosa (Villalard-Bohnsack and Harlin 1997), while in Brittany G. turuturu was overgrown with P. fibrata and Ceramium rubrum (Simon et al. 2001). However, in Narragansett Bay and Long Island Sound, G. turuturu had a low biomass, density, and diversity compared to native Chondrus crispus (Janiak and Whitlach 2012; Jones and Thornber 2010). In Narragansett Bay, epiphytes of G. turuturu were frequently grazed by the small snail Lacuna vincta (Jones and Thornber 2010). However, in Long Island Sound, the overall abundance and diversity of epifaunal invertebrates on G. turuturu was low compared to that on Chondrus crispus (Janiak and Whitlach 2012). We have found no mention of direct grazing on G. turuturu.

Trophic Status:

Primary Producer

PrimProd

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Habitats

General Habitat	Rocky	None
General Habitat	Oyster Reef	None
General Habitat	Marinas & Docks	None
General Habitat	Vessel Hull	None
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

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Life History

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	3	Field (Narragansett Bay, Harlin and Villaland-Bohnsack 2001)
Maximum Temperature (ºC)	28	Experimental (Brittany, Simon et al. 1999; Wei et al. 2013). Plants died at 30 C (Wei er al. 2013).
Minimum Salinity (‰)	15	Field (Simon et al. 1999). Experimentally, photosynthesis was slightly reduced at 22 PSU, but sharply at 12 PSU (Simon et al. 1999).
Maximum Salinity (‰)	42	Experimentally, photosynthesis was slightly reduced at 42 PSU, but sharply at 52 PSU (Simon et al. 1999).
Maximum Length (mm)	2,950	Brittany, France (Simon et al. 2001), more usually to 900-1000 mm (Simon et al. 2001; Van Patten 2006)
Broad Temperature Range	None	Cold temperate-Warm temperate
Broad Salinity Range	None	Polyhaline-Euhaline

General Impacts

To our knowledge, there are no known economic impacts of Grateloupia turuturu and ecological impacts appear to be localized.

Competition- Qualitative observations in Mexico and France suggest that G. turuturu can compete with and replace native seaweeds (Simon et al. 2001; Aguilar-Rosas et al. 2012). In Brittany, growth of G. turuturu appeared to be stimulated by eutrophication (Simon et al. 2012). In Portugal, removal experiments suggest that G. turuturu is an opportunist, favored by disturbances which remove natives and therefore a ‘passenger’ rather than a ‘driver’ of ecological changes (Mulas and Bertocci 2016). Its growth in rockpools was enhanced by a combination of disturbance and nutrient enrichment (Bertocci et al. 2014).

Habitat Change- Observations on the East Coast, in Long Island Sound and Narragansett Bay, suggest that Grateloupia turuturu supports fewer epiphytes and fewer invertebrates (in Long Island Sound) than the native red alga Chondrus crispus (Jones and Thornber 2010; Janiak and Whitlach 2012). In Narragansett Bay, biomass and diversity of epiphyytes were low on G. turuturu and the native brown alga Fucus vesiculosus, compared to C. crispus and the introduced green alga Codium fraglie (Jones and Thornber 2010). In Long Island Sound, only C. crispus and G. turuturu were compared. Both epiphytes and epifaunal invertebrates were found to have lower density and diversity on G. turuturu, with the implication that the spread of G. turuturu could reduce biodiversity in invaded habitats (Janiak and Whitlach 2012). It should be noted that while the diversity of epiphytes was lower on G. turuturu, it still supported 24 species of epiphytes in Narragansett Bay, including the introduced red algae Antithamnion hubbsi and Neosiphonia harveyi (Jones and Thornber 2010). 

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Regional Impacts

NA-ET3	Cape Cod to Cape Hatteras		Ecological Impact	Habitat Change
Shallow subtidal algal communities at Millstone Point, CT, dominated by G. turuturu had reduced epiphyte biomass and reduced abundance and species richness of invertebrates (Janiak and Whitlach 2012).
M040	Long Island Sound		Ecological Impact	Habitat Change
Shallow subtidal algal communities at Millstone Point, CT, dominated by G. turuturu had reduced epiphyte biomass and reduced abundance and species richness of invertebrates (Janiak and Whitlach 2012).
NEP-VI	Pt. Conception to Southern Baja California		Ecological Impact	Competition
Replacing native species in Ensenada, Mexico (Aguliar-Rosas et al. 2012; Aguliar-Rosas et al. 2014)
NEA-V	None		Ecological Impact	Competition
In experiments at Foz do Douro, Portugal, G. turuturu was most abundant in pools where native canopy-forming seaweeds were continuously removed, compared to controls. In pools where G. turuturu was continuously removed, overall community composition did not differ from controls, suggesting that G. turuturu is a 'passenger' of disturbance, rather than a driver of community change (Mulas and Bertocci 2016). However, individual species responded in idiosyncratic ways, varying with habitat (tidepools vs. exposed rock) and shore level. Among species showng these differens were: Chondrus crispus; articulated coralline algae, Corallina spp.; the crustose sporophyte of the red alga Mastocarpus stellatus, and the mussel, Mytilus galloprovincialis (Freitas et al. 2016). Nutrient fertilization of tidepools promoted the establishment and functional impacts (increased productivity and respiration) by Grateloupia turuturu and Sargassum muticum (Vieira et al. 2017).

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