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

---

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).

---

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.

---

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

---

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

---

---

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). 

---

Regional Impacts

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)

References

Aguilar-Rosas, Luis Ernesto; Pedroche, Francisco Flores; Zertuche-González, José Antonio (2014) [Aquatic Invasive Species in Mexico], Comisión Nacional para el Conocimiento y Uso de la Biodiversidad, <missing place>. Pp. 211-222

Aguilar-Rosas,Luis E.; Boo, Sung Min; Kim, Kyeong Mi; Aguilar-Rosas, Cristiane V. (2012) [First record of the Japanese species Grateloupia turuturu (Halymeniaceae, Rhodophyta) in the Mexican Pacific coast], Hidrobiológica 22(2): 189-194

Araujo, Rita and 5 authors 1*, JOSE´ VIOLANTE 1, RUI PEREIRA 1, HELENA ABREU 1, FRANCISCO ARENAS 1 AND ISABEL SOUSA-PINTO (2011) Distribution and population dynamics of the introduced seaweed Grateloupia turuturu (Halymeniaceae, Rhodophyta) along the Portuguese coast, Phycologia 50(4): 392-402

Ardré, Françoise, Gayral, Paulette (1961) Quelques Grateloupia de l'Atlantique et du Pacifique, Revue Algologique 6: 38-48

Arenas, F. and 13 authors. (2006) Alien species and other notable records from a rapid assessment survey of marinas on the south coast of England., Journal of the Marine Biological Association of the United Kingdom 86: 329-1337

Bárbara, Ignacio; Cremades, Javier (2004) Grateloupia lanceola versus Grateloupia turuturu (Gigartinales, Rhodophyta) en la Península Ibérica, Anales del Jardin Botanico de Madrid 61(2): 103-118

Barreiro, Rudolfo; Quintela, Maria, Barbara, Ignacio; Cemades, Barbara (2006) RAPD differentiation of Grateloupia lanceola and the invasive Grateloupia turuturu (Gigartinales, Rhodophyta) on the Iberian Peninsula., Phycologia 45(2): 213-217

Bertocci, Iacopo and 9 authors (2014) The regime of climate-related disturbance and nutrient enrichment modulate macroalgal invasions in rockpools, Biological Invasions Published online: <missing location>

Bold, Harold C.; Wynne, Michael J. (1978) Introduction to the Algae: Structure and Reproduction, Prentice-Hall, Englewood Cliffs, NJ. Pp. <missing location>

California Department of Fish and Wildlife (2014) Introduced Aquatic Species in California Bays and Harbors, 2011 Survey, California Department of Fish and Wildlife, Sacramento CA. Pp. 1-36

Carreira-Flores, Diego; Rubal, Marcos; Moreira, Juan; Guerrero-Meseguer, Laura; Gomes, Pedro T.; Veiga, Puri (2023) Recent changes on the abundance and distribution of non-indigenous macroalgae along the southwest coast of the Bay of Biscay, Aquatic Botany 189(103685): Published online
https://doi.org/10.1016/j.aquabot.2023.103685

Cecere, Ester; Moro, Isabella; Wolf, Marion A.; Petrocelli, Antonella; Verlaque, Marc; Sfriso, Adriano (2011) The introduced seaweed Grateloupia turuturu (Rhodophyta, Halymeniales) in two Mediterranean transitional water systems, Botanica Marina 54: 23-33

Chainho, Paula and 20 additional authors (2015) Non-indigenous species in Portuguese coastal areas, lagoons, estuaries, and islands, Estuarine, Coastal and Shelf Science <missing volume>: <missing location>

Çinar, Melih Ertan and 7 authors (2021) Current status (as of end of 2020) of marine alien species in Turkey, PLOS ONE 16: Published online

D'Archino, R. D.; Nelson, W. A.; Zuccarello, G. C. (2007) Invasive marine red alga introduced to New Zealand waters: first record of Grateloupia turuturu (Halymeniaceae, Rhodophyta)., New Zealand Journal of Marine and Freshwater Research 41: 35-42

de Azevedo, Carolina Angelica Araujo; Cassano, Valeria; Horta, Paulo Antunes Junior; Batista, Manuela Bernardes; de Oliveira, Mariana Cabral (2015) Detecting the non-native Grateloupia turuturu (Halymeniales, Rhodophyta) in southern Brazil, Phycologia 54(5): 451-454

Eno, N. Clare; Clark, Robin A.; Sanderson, William G. (1997) <missing title>, Joint Nature Conservation Committee, Peterborough. Pp. <missing location>

Farnham, W. F. (1980) Studies on aliens in the marine flora of southern England, In: Price, J. H., Irvine, D. E. G., and Farnham, W. F.(Eds.) The Shore Environment. , London. Pp. 875-914

Farnham, William F. (1994) Introduction of marine benthic algae into Atlantic European waters, In: Boudouresque, C. F., Briand, F., and Nolan, C.(Eds.) Introduced Species in European Coastal Waters. , Brussels. Pp. 32-36

Fernandez-Alvarez, Fernando Angel Machordom, Annie (2012) DNA barcoding reveals a cryptic nemertean invasion in Atlantic and Mediterranean waters, Helgoland Marine Research 67: 599-605
DOI 10.1007/s10152-013-0346-3

Ferreira, S.; Kaufmann, M.; Neto, A.; Izaguirre,, J. P.; Wirtz, P.; De Clerck. O. 2012 New records of macroalgae from Madeira Archipelago. <missing URL>



Gabriel, Daniela; Ferreira, Ana Isabel; Micael, Joana; Fredericq, Suzanne (2023) Non-Native Marine Macroalgae of the Azores: An Updated Inventory, Diversity 15(1089): Published online
https://doi.org/10.3390/ d15101089

Gavio, Brigitte; Fredericq, Suzanne (2002) Grateloupia turuturu (Halymeniaceae, Rhodophyta) is the correct name of the non-native species in the Atlantic known as Grateloupia doryphora., European Journal of Phycology 37(3): 349-359

Gittenberger, Arjan; Mirimin, Luca; Boyd, John; O’Beirn, Francis; Devine, Grainne; O’Brien, Martina; Rensing, Marjolein; O’Dwyer, Katie; Gittenberger, Edmund (2023) Marine Non-Indigenous Species Dynamics in Time and Space within the Coastal Waters of the Republic of Ireland, Diversity 15(1019): Published online
https:// doi.org/10.3390/d15091019

Gorostiaga, Jose M.; Santolaria, Alberto; Secilla, Antonio; Casares, Coro; Diez, Isabel (2004) Check-list of the Basque coast benthic algae (North of Spain)., Anales del Jardin Botanico de Madrid 61(2): 155-180

Goulletquer, Philippe; Bachelet, Guy; Sauriau, Pierre; Noel, Pierre (2002) Invasive aquatic species of Europe: Distribution, impacts, and management, Kluwer Academic Publishers, Dordrecht. Pp. 276-290

Gravili, Cinzia and 12 authors (2010) Nonindigenous species along the Apulian coast, Italy, Chemistry and Ecology 26: 121-142

Guidone, Michele; Thornber, Carol; Brian Wysor, Brian Wysor; O’Kelly, Charles J. (2013) Molecular and morphological diversity of Narragansett Bay (RI, USA) Ulva (Ulvales, Chlorophyta) populations, Journal of Phycology 49: 979-995
DOI: 10.1111/jpy.12108

Guiry, M. D.; Guiry, G. M. 2004-2023 AlgaeBase. https://www.algaebase.org/



Harlin, M. M.; Villalard, Bohnsack (2001) Seasonal dynamics and recruitment strategies of the invasive seaweed Grateloupe doryphora (Halymeniaceae, Rhodophyta) in Narragansett Bay and Rhode Island Sound, RI, USA, Phycologia 40(5): 468-474

Harlin, Marilyn M., Villalard-Bohnsack, Martine (1999) A large red seaweed invades Narragansett Bay, Maritimes 41(4): 6-9

Hughey, Jeffery R.; Miller, Kathy Ann; Lyman, Ashleigh (2009) Noteworthy Collections: California, Madroño 56(4): 293-295

Inderjit; Chapman, David ; Ranelletti, Marla ; Kaushik, Shalini (2006) Invasive marine algae: an ecological perspective., Botanical Review 72(2): 153-178

Janiak, Dean S.; Whitlatch, Robert B. (2012) Epifaunal and algal assemblages associated with the native Chondrus crispus (Stackhouse) and the non-native Grateloupia turuturu (Yamada) in eastern Long Island Sound, Journal of Experimental Marine Biology and Ecology 413: 38-44

Jones, Emily; Thornber, Carol S. (2005) Effects of habitat-modifying invasive macroalgae on epiphytic algal communities, Marine Ecology Progress Series 400: 87-100

Katsanevakis, S. and 39 authors (2014) New Mediterranean Biodiversity Records (October, 2014), Mediterranean Marine Science 15(3): 675-695

Kennedy, C., Pappal, A. L.; Bastidas, C.; ; Carlton, J. T.; David, A. A.; Dijkstra, J.A.; Duffey, S; Gibson, J.; Grady, S. P.; Green-Gavrielidis, (2020) Report on the 2018 Rapid Assessment Survey of Introduced, Cryptogenic, and Native Marine Species at New England Marinas: Massachusetts to Maine, <missing publisher>, Boston MA. Pp. <missing location>

Kinoshita, Kyoko; Nakayama; Satoko; Furota, Toshio (2003) Life cycle characteristics of the deep-burrowing mud shrimp Upogebia major (thalassinidea: upogebiidae)on a tidal flat along the northern coast of Tokyo Bay, Journal of Crustacean Biology 23(2): 318-327

Lee, Ju Il; Kim, Hyung Geun; Geraldino, Paul John L; Hwang, Il Ki; Boo, Sung Min (2009) Molecular classification of the genus Grateloupia (Halymeniaceae, Rhodophyta) in Korea, Algae 24(4): 231-238

Liu, Wenliang; Liang, Xiaoli ; Zhu, Xiaojing (2015) A new record and mitochondrial identification of Synidotea laticauda Benedict, 1897 (Crustacea: Isopoda: Valvifera: Idoteidae) from the Yangtze Estuary, China, Zootaxa 4294: 371-380

Lyons, Devin A.; Scheibling, Robert E. (2009) Range expansion by invasive marine algae: rates and patterns of spread at a regional scale, Diversity and Distributions 15: 762-775

MacIntyre, Chris; Adrienne Pappal; Pederson, Judy; Smith, Jan P. (2011) Marine Invaders in the Northeast Rapid Assessment Survey of non-native and native marine species of floating dock communities, Massachusetts Coastal Zone Management, Boston MA. Pp. <missing location>

Maggs, Christine A.; Stegenga, Herre (1999) Red algal exotics on North Sea coasts., Helgoländer Meeresuntersuchungen 52: 243-258

Marston, Marcia; Villaland-Bohnsack, Martine (2002) Genetic variability and potential sources of Grateloupia doryphora, an invasive species in Rhode Island waters., Journal of Phycology 38: 649-658

Marston, Marcia; Villalard-Bohnsack, Martine (2000) The use of molecular genetics to investigate the geographical origin and vector of an invasive red alga., In: Pederson, Judith(Eds.) Marine Bioinvasions. , Cambridge. Pp. 244-250

Massachusetts Office of Coastal Zone Management (2013) Rapid assessment survey of marine species at New England floating docks and rocky shores, Massachusetts Office of Coastal Zone Management, Boston MA. Pp. <missing location>

Mathieson, A. C. (1992) Transport of seaweeds by tagged migratory fish populations, Botanica Marina 35: 419-422

Mathieson, Arthur C.; Dawes, Clinton J. (2017) Seaweeds of the Northwest Atlantic, University of Massachusetts Press, Amherst MA. Pp. <missing location>

Mathieson, Arthur C.; Dawes, Clinton J.; Pederson, Judith; Gladych, Rebecca; Carlton, James T. (2007) The Asian red seaweed Grateloupia turuturu (Rhodophyta) invades the Gulf of Maine., Biological Invasions (preprint): <missing location>

Mathieson, Arthur C.; Pederson, Judith R.; Neefus, Christopher D.; Dawes, Clinton J.; Bray, Troy L. (2008) Multiple assessments of introduced seaweeds in the Northwest Atlantic., ICES Journal of Marine Science 65: 730-741

Micael, Joana; Parente, Manuela I.; Costa, Ana C. (2014) Tracking macroalgae introductions in North Atlantic oceanic islands, Helgoland Marine Research 68: 209-219

Miller, Kathy Ann 2016 Seaweeds of California. <missing URL>



Miller, Kathy Ann; Aguilar-Rosas, Luis Ernesto; Pedroche, Francisco F. (2011) A review of non-native seaweeds from California, USA and Baja California, Mexico, Hidrobiológica 21(3): 365-379

MIT Sea Grant 2003-2008 Introduced and cryptogenic species of the North Atlantic. <missing URL>



MIT Sea Grant 2009-2012 Marine Invader Tracking and Information System (MITIS). <missing URL>



Mulas, Martina; Bertocci, Iacopo (2016) Devil's tongue weed (Grateloupia turuturu Yamada) in northern Portugal: Passenger or driver of change in native biodiversity?, Marine Environmental Research 118: 1-9

Nyberg, Cecilia D.; Wallentinus, Inger (2005) Can species traits be used to predict marine macroalgal introductions?, Biological Invasions 7: 265-279

O’Shaughnessy, Kathryn A.; Harding, Juliana M.; Burge, Erin J. (2014) Ecological effects of the invasive parasite Loxothylacus panopaei on the flatback mud crab Eurypanopeus depressus with implications for estuarine communities, Bulletin of Marine Science 90(2): 611-621

Oliveira, Otto M. P. and 24 authors (2016) Census of Cnidaria (Medusozoa) and Ctenophora from South American marine waters, Zootaxa 4194: 1-256

Pederson, Judith, and 13 authors (2021) 2019 Rapid Assessment Survey of marine bioinvasions of southern New England and New York, USA, with an overview of new records and range expansions, Bioinvasions Records 10(2): 22-–237

Peña, V.; Bárbara, I.; Grall, J.; Maggs, C. A.; Hall-Spencer, J. M. (2014) The diversity of seaweeds on maerl in the NE Atlantic, Marine Biodiversity 44: 533-551

Petrocelli, Antonella; Carlucci, Roberto; Portacci, Giuseppe; Cecere, Ester (2011) Phenology of the alien seaweed Grateloupia turuturu population in the Mar Piccolo of Taranto, Journal of Phycology 46(Supplement 1): 184-185

Plouguerne, Erwan; Hellio, Claire; Deslandes, Eric; Veron, Benoit; Stiger-Pouvreau, Vale rie (2008) Anti-microfouling activities in extracts of two invasive algae: Grateloupia turuturu and Sargassum muticum., Botanica Marina 51: 202-208

Ruiz, Gregory M.; Geller, Jonathan (2018) Spatial and temporal analysis of marine invasions in California, Part II: Humboldt Bay, Marina del Re, Port Hueneme, and San Francisco Bay, Smithsonian Environmental Research Center & Moss Landing Laboratories, Edgewater MD, Moss Landing CA. Pp. <missing location>

Ryu, Tae-Bok; Choi, Dong-Hui; Kim, Deokki; Lee, Jung-Hyo; Lee, Do-Hun; Kim, Nam-Young (2018) Distribution and current vegetation of Cakile edentula, an invasive alien species in Korea, Journal of Ecology and Environment 43(15): Publkished online
https://doi.org/10.1186/s41610-018-0072-5

Saunders, Gary W.; Withall, Rodney D. (2006) Collections of the invasive species Grateloupia turuturu (Halymeniales, Rhodophyta) from Tasmania, Australia., Phycologia 45(6): 711-714

Simon, C.; Ar Gall, E. ; Levavasseub, G.; Deslandes, E. (1999) Effects of short-term variations of salinity and temperature on the photosynthetic response of the red alga Grateloupia doryphora from Brittany (France), Botanica Marina 42: 437-440

Simon, C.; Gall, A. R.; Deslandes, E. (2001) Expansion of the red alga Grateloupia doryphora along the coasts of Brittany, Hydrobiolgia 443: 23-29

Soors, Jan; Faasse, Marco; Stevens, Maarten; Verbessem, Ingrid; De Regge, Nico;Van den Bergh, Ericia (2010) New crustacean invaders in the Schelde estuary (Belgium), Belgian Journal of Zoology 140: 3-10

Stegenga, Herre; Karremans, Mart (2015) [Review of the red algal exotics in the marine waters of the Southwest Netherlands], Gorteria 27: 141-157

Stiger-Pouvreau, Valérie; Thouzeau, Gérard (2015) Marine species introduced on the French Channel-Atlantic coasts: a review of main biological invasions and impacts, Open Journal of Ecology, 5: 227-257

Stuckey, F. G. (1909) Notes on a New Zealand actinian, Bunodes aureoradiata, Transactions and Proceedings of the New Zealand Institute 41: 367-369

Tagliapietra, Davide; Guarneri , Irene; Keppel, Erica; Sigovini, Marco (2021) After a century in the Mediterranean, the warm-water shipworm Teredo bartschi invades the Lagoon of Venice Italy), overwintering a few degrees above zero, Biological Invasions 23: 1595-1618

Texeira, Valeria L., Almeida, Sergio A. D. S., Kelecom, Alphonse (1990) Chemosystematics and biogeographic studies of the diterpenes from the marine brown alga Dictyota dichotoma, Biochemical Systematics and Ecology 18(2/3): 87-92



Van Patten, Margaret Stewart (2006) Seaweeds of Long Island Sound, Connecticut Sea Grant, Groton. Pp. <missing location>

Vázquez-López, Horacio; Cházaro-Olvera, Sergio; Vargas-Téllez, Irma ; Molina-Ortega, Madeline Getzemany (2017) Description of first zoeal stage of Cardisoma crassum Smith, 1870 (Crustacea: Decapoda: Gecarcinidae), Journal of Natural History 51(15-19): 837-842

Verlaque, Marc (2001) Checklist of the macroalgae of Thau Lagoon (Herault, France), a hot spot of marine species introduction in Europe, Oceanologia Acta 24(1): 29-49

Verlaque, Marc; Brannock, Pamela M.; Komatsu, Teruhisa; Villalard-Bohnsack, Martine; Marston, Marcia (2005) The genus Grateloupia (Halymeniaceae, Rhodophyta) in the Thau Lagoon (France, Mediterranean); a case study of marine plurispecific introductions., Phycologia 44(5): 477-496

Villalard-Bohnsack, Martine, Harlin, Marilyn M. (1997) Appearance of Grateloupia doryphora (Halymeniaceae, Rhodophyta) on the northeast coast of North America, Phycologia 36(4): 324-328

Villalard-Bohnsack, Martine, Harlin, Marilyn M. (1997) An invasive red seaweed enters Narragansett Bay, RINHewS 4(1): 2-3

Wallentinus, Inger 2002 Introduced marine algae and vascular plants in European aquatic environments.. <missing URL>



Wallentinus, Inger (2002) Introduced marine algse and vascular plants in European aquatic environments., In: Leppakoski, E.; Gollasch, S.; Olenin, S.(Eds.) Invasive aquatic species of Europe: distribution, impacts and management. , Dordrecht/ Boston/ London. Pp. 27-52

Wei, Xiaojiao; Shuai, Limei; Lu, Bojun; Wang, Shasha; Chen, Jia; Wang, Gaoge (2013) Effects of temperature and irradiance on filament development of Grateloupia turuturu (Halymeniaceae, Rhodophyta), Journal of Applied Phycology 25: 1881-1886

Wells, Christopher D. and 23 authors (2014) Report on the 2013 rapid assessment survey of marine species at New England bays and harbors, Massachusetts Office of Coastal Zone Management, Boston MA. Pp. 32

Yang, Mi Yeon; Kim, Myung Sook (2015) Taxonomy of Grateloupia (Halymeniale, Rhodophyta) by RNA barcode marker analysis and a description of Pachymeniopsis volvita sp. Nov., Journal of Applied Phycology 27: 1373-1384

Zabin, Chela; Guerra, Vanessa; Grosholz, Edwin D.; Lesyna, Kristine; McGowan,Jennifer (2011) <missing title>, San Francisco Regional Water Quality Control Board, San Francisco. Pp. 31

---