Codium fragile ssp. Fragile

Invasion History

First Non-native North American Tidal Record: 1957
First Non-native West Coast Tidal Record: 1977
First Non-native East/Gulf Coast Tidal Record: 1957

---

General Invasion History:

Codium fragile is believed to have originated in the Northwest Pacific in waters around Japan. In Europe, it was first collected on the west coast of Ireland in 1845, but not recognized as an introduced species until around 1900 (Chapman 1999; Provan et al. 2008). A morphotype, C. f. subsp. atlanticum, of uncertain taxonomic and invasion status, was collected even earlier in 1839, in Ireland, but has had a limited range in Europe (Provan et al. 2008). The more typical C. fragile was first collected in the Mediterranean in 1950 (Verlaque 1994). In North America, Codium fragile was first collected on Long Island, New York in 1957 (Bouck and Morgan 1957). This seaweed spread rapidly to the north and south, reaching Nova Scotia in 1991 (Begin and Scheibling 2003); the Magdalen Islands, in the Gulf of St. Lawrence, Quebec, by 2003 (Drouin et al. 2011); and the south coast of Newfoundland by 2012 (Matheson et al. 2014). To the south, it colonized Beaufort Inlet, North Carolina in 1981 (Searles et al. 1984) and the current southernmost record is Cape Fear Inlet, North Carolina (Geraldi et al. 2014).

On the West Coast, at least one separate species of Codium or a separate subspecies of C. fragile is native from Baja California to Alaska, but an introduced population of C. fragile was discovered in San Francisco Bay in 1977 (Silva 1979; Cohen and Carlton 1995). Other invasions by C. fragile are widely scattered in space and time, including the Falkland Islands in 1849 (Provan et al. 2008); Cape Horn in1842; New Zealand in 1841 (Provan et al. 2008); Cape Town, South Africa in 1884 (Provan et al. 2008); Tasmania in 1890 (Provan et al. 2008); Iceland, before 1985 (South and Tittley 1986); Azores in 1993 (Cardigos et al. 2006); and Chile in 1998 (Castilla et al. 2005). Some of these populations were named as new subspecies, but were found, by genetic methods, to belong to C. fragile subsp. fragile (Provan et al. 2008).

North American Invasion History:

Invasion History on the West Coast:

Early Codium specimens from the West Coast of North America were identified as C. mucronatum forma californicum ( J. Agardh 1887), later known as C. fragile subsp. californicum (J. Agardh) C.A. Maggs & J. Kelly (Guiry and Guiry 2016). This form of Codium is found from Baja California to Alaska, mostly on the open coast (Abbott and Hollenberg 1976; Silva 1979; Miller et al. 2011). In 1977, a population of C. fragile, then identified as subspecies tomentosoides, was found only at Coyote Point in San Francisco Bay (Silva 1979). Codium fragile was found at Coyote Point and Redwood City in 1978-1983 (Josselyn and West 1985). In 1993-94, Cohen and Carlton (1995) found it growing on floating docks from Richmond to San Leandro, and a Pier 39, in San Francisco Harbor (Cohen and Carlton 1995). This seaweed has also been collected in Tomales Bay (Miller 2004; Miller et al. 2011).

Invasion History on the East Coast:

Codium fragile's first appearance in Northwest Atlantic waters was in Gardiners Bay, Long Island in 1957 (Bouck and Morgan 1957). Its introduction has been attributed to ship fouling, ballast water, or to the introduction of European or Japanese oysters. The first mode seems likeliest, because precautions were taken to prevent introduction of fouling organisms with European oysters planted experimentally in 1949 (Loosanoff 1975; cited by Carlton and Scanlon 1985) and because the dates and sites of oyster plantings do not correspond with those of C. fragile. The spotty nature of its spread in East coast waters suggests a diversity of transport mechanisms. Codium fragile spp. fragile was first collected in East Marion New York on the Eastern Fork of Long Island, in Gardiners's and Peconic Bays in 1957 (Bouck and Morgan 1957). It reached the Connecticut side of Long Island Sound by 1961 and Rhode Island by 1962-68. It was then brought to Chatham and Cotuit on Cape Cod in 1961 with transplanted oysters from Long Island. By the early 70's, it was widespread on the south side of Cape Cod (Carlton and Scanlon 1985).

Codium fragile was first collected north of Cape Cod in Boothbay Harbor Maine in 1964, where it was probably introduced with transplanted oysters (Carlton and Scanlon 1985). Its spread in Cape Cod Bay occurred later. It reached the Cape Cod Canal by 1969, and successively was found in Barnstable Harbor, MA in 1972; Wellfleet Harbor, MA in 1974; Duxbury Bay in 1981; and Provincetown in 1981. By 1983, it was collected at Appledore Island, Isles of Shoals, Maine (Carlton and Scanlon 1985). However, it was present to the north, at Bar Harbor, Maine by the late 1970's or early 1980's (Bird et al. 1993). The northward spread was spotty, with first records in Massachusetts Bay (Salem) in 2000 (MIT Sea Grant 2003), and in Great Bay, New Hampshire in 1985 (Mathieson et al. 2003). Codium fragile was first collected in Mahone Bay, on the central Atlantic coast of Nova Scotia in 1991. It has not yet been seen in the Bay of Fundy, although it was found in Sam Orr's Pond, a tributary of Passamaquoddy Bay, New Brunswick in 2009 (Saunders et al. 2013). It may have been brought to Nova Scotia, with transplanted shellfish, as packing material for bait, or by Gulf Stream eddies from New England (Bird et al. 1993).

In 1996, Codium fragile was collected for the first time in the Gulf of St. Lawrence, in Caribou Harbour, on the Northumberland Straits (Garbary et al. 1997). Since then, the spread has continued, both in the Gulf of St. Lawrence, and on the Atlantic Coast of Nova Scotia and Newfoundland. Codium fragile has been spreading through Northumberland Straits, in Nova Scotia, New Brunswick and on the Straits and Gulf Side of Prince Edward Island (Hubbard and Garbary 2002; Garbary et al. 2004). One population in Malpeque Bay included plants with the morphology of C. f. subsp. atlanticum (Hubbard and Garbary 2002). In the Gulf, it has spread as far north as the Magdalen Islands, Quebec in 2002 (Simard et al. 2005; Drouin et al. 2011, 47.3 N). Codium fragile is now patchily distributed on the whole Atlantic Coast of Nova Scotia, from Cape Sable Island, at the southwestern end to Canso, a distance of 450 km (Hubbard and Garbary 2002; Watanabe et al. 2010). In 2012, C. fragile was found at several sites in Placentia Bay, Newfoundland (Matheson et al. 2014, 47.8 N).

Between 1957 and 1968, C. fragile became widespread and abundant in Long Island Sound (Carlton and Scanlon 1985). It was first found in Barnegat Bay in 1966 and was widely distributed by 1969 (Taylor et al. 1969). The next southern record was in Assateague Channel, part of Chingcoteague Bay, Virginia in 1976 (Hillson 1976). Surprisingly, C. fragile was not collected at other estuaries in the region until much later. It was found washed up on the beach in Poquoson Neck, Virginia, on the western side of the lower Chesapeake Bay in 1995 (Scott Godwin, personal communication). It was found at Cape Henry, Virginia in 1999 (Ruiz et al., unpublished data), and was found in Atlantic coastal bays from Indian River Bay, Delaware to Hog Island Bay, Virginia in 2000-2003 (Thomsen 2004; Miller and Brown 2005; Thomsen and McGlathery 2006). However, we have no records from Delaware Bay. Codium fragile approached its southernmost limit at Topsail Inlet, south of Cape Hatteras, North Carolina in 1979, Masonboro Inlet in 1982, and Beaufort Inlet in 1981 (Searles et al. 1984). It was found growing abundantly on artificial substrates in Cape Fear Sound, near Wilmington, North Carolina in 2009 (Geraldi et al. 2014).

Invasion History Elsewhere in the World:

Codium fragile was an early invader to European waters, but was unrecognized. The possibly synonymous or related C. f. subsp. atlanticum was collected in Ireland in 1839 (Provan et al. 2008). However, taxonomists disagree on whether this form is introduced or native in Europe (Silva 1955; Provan et al. 2008; Guiry and Guiry 2016). It has a mostly European distribution from Norway to Spain, but 'atlanticum-like' plants have been found in Nova Scotia and the Azores (Hubbard and Garbary 2002; Tittley and Neto 2005). The typical C. fragile was collected on the west coast of Ireland in 1845, but was mistaken for the native C. tomentosum (Provan et al. 2008). It and other early C. fragile herbarium specimens were re-identified using chloroplast DNA (Provan et al. 2008). Codium fragile was recognized as a non-native species when it was collected in the Netherlands in 1900 (Silva 1955). However, it had already been collect in Devon, England in 1853 (Provan et al. 2008). It has spread extensively along the coast of Europe, reaching Sweden by 1939, the Danish Kattegatt (Baltic mouth) by 1985 (Chapman 1999), and Galicia, Spain by 1989 (Pérez-Cirera et al. 1989). It was first collected in the Mediterranean in France in 1950 (Verlaque 1994) and has spread east, reaching the Turkish Aegean by 1985 (Ribera and Boudouresque 1995). In the eastern Atlantic, C. fragile was established in Iceland by 1985 (South and Tittley 1987), the Canary Islands by 1990 (Haroun et al. 2002) and the Azores by 1993 (Tittley and Neto 2005).

Codium fragile was introduced to several locations in the Southern Hemisphere in the 18th century, including Cape Horn, Chile in 1842; the Falkland Islands in 1849; Bay of Islands, New Zealand in 1841; and the Cape of Good Hope, South Africa in 1884 (Provan et al. 2008). We have not found information on established populations in Atlantic South America, but C. fragile is established in several locations on the northern and central coasts of Chile (Castilla et al. 2005; Neill et al. 2006). Codium fragile is established on the southern Atlantic coast of South Africa, but its range is unclear, because of confusion with presumed native forms of Codium (Mead et al. 2011b). The South African form was described as C. f. capensis, but the type specimen proved to be genetically identical to C. fragile (Provan et al. 2008). Similarly, in New Zealand, a native subspecies, C. f. nova-zelandiae was described in 1935, but the type specimen was found to actually be C. fragile (Provan et al. 2008). Prior to this discovery, the introduced C. fragile was thought to have been introduced to New Zealand in 1973 and to have a limited range near Auckand (Dromgoole and Foster 1983; Trowbridge 1995). In Australia, Codium fragile subsp. tasmanicum was described from Tasmania in 1889, but was also found to be identical to typical C. fragile (Provan et al. 2008). A herbarium specimen of C. fragile was collected in Port Philip Bay in 1890 (Provan et al. 2008), but the first reported discovery of this plant was made in 1996 (Campbell 1999). It was subsequently found near Sydney in 2001 (NIMPIS 2015); in West Lakes, South Australia in 2002 (Wiltshire et al. 2010); and Albany, Western Australia in 2008 (McDonald et al. 2015).

---

Description

Codium fragile has thick, spongy, finger-like and forking branches, which extend from an irregular holdfast. The branches are covered with densely packed, short hairs, giving the plant a felt-like appearance and slimy texture. The tips of the branches are tapered. A plant may be about 0.5-1 m tall, with about 12 orders of branching, and weigh up to 3 kg. The branches are buoyant in the water due to the production of oxygen inside the plant. Older branches are dark green, while the tips and newer branches are a bright lighter green. Dead branches are whitish-yellow and resemble thick spaghetti. This species has many descriptive common names, including 'Dead-Man's Fingers', 'Green Fleece', Sponge Weed', 'Sponge Fingers', ‘Sputnik Weed’, 'Staghorn Weed' and 'Oyster Thief'. Oyster Thief describes the tendency of the oxygen-filled weeds to float and carry away oysters, while Sputnik Weed refers to its mysterious appearance around Long Island in 1957, soon after the launching of the Russian satellite.

The body of a green alga is called a thallus, a mass of undifferentiated filaments ('siphons'). In the genus Codium, the thallus is thick and spongy, and formed into the distinctive dichotomous branches. The internal structure of the thallus is comprised of a medulla, densely packed colorless siphons, surrounded by a green palisade-like layer of vesicles called utricles. The siphons are coenocytic and lack walls between the cells. The uitricles are the outer tips of these filaments. The holdfast is composed of a web of rhizoids (root-like filaments). The utricles in C. fragile are club-shaped, with a small pointed tip. Utricles bear one or two hairs below the tip, 25-60 µm long, 50-600 µm long. The dense field of hairs gives the branches a fuzzy appearance. The growth form is highly variable. This description is based on: Silva 1955, Bold and Wynne 1978, Gosner 1978, Schneider and Searles 1991, Trowbridge 1998, Van Patten 2006, Fisheries and Oceans Canada 2011, and Guiry and Guiry 2016.

The genus Codium is challenging taxonomically. At least 130 species are recognized, mostly in tropical and subtropical zones (Appeltans et al. 2016). For example, the occurrence of C. fragile in Europe was overlooked for at least 50 years after its introduction, because of its similarity to the native European C. tomentosum (Provan et al. 2008). The taxonomy of C. fragile is also complex at the subspecies level. At least 17 subspecies have been recognized, from different parts of the world, but the status of these is uncertain. Codium fragile was originally described by Suringa, from specimens collected in Japan, in a book by Hariot et al. 1889 (cited by Provan et al. 2008). European specimens were identified as Codium fragile subspecies tomentosoides, based on the type from the Netherlands, collected in 1900 (Silva 1955). This name was widely applied to introduced populations in many parts of the world, including the East Coast of North America (Bouck and Morgan 1957), San Francisco Bay (Silva 1979), Australia (Campbell 1999), and New Zealand (Trowbridge 1998). A worldwide genetic survey of the subspecies of C. fragile concluded that the invasive C. f. subsp. tomentosoides was identical to the original type form, and should be known as C. f. subsp. fragile, or simply C. fragile (Provan et al. 2008; Guiry and Guiry 2016).

Specimens of three named geographical subspecies, including subsp. capensis from South Africa, subsp. novae-zelandiae from New Zealand, and subsp. tasmanicum proved to be C. fragile (Provan et al. 2008). One or more apparently native Codium species grow on the West Coast of North America from Alaska to Mexico, with a sporadic distribution on rocks of the outer coast. The apparently native plants have been known as C. f. subsp. californicum, and appear to be distinct from the introduced C. fragile growing in San Francisco Bay (Silva 1979; Trowbridge 1998; Miller et al. 2011). However, Provan et al. (2009) found that one herbarium specimen, marked as C. f. subsp. californicum from Washington State belonged with the other C. fragile populations. Armitage et al. (2017) did find that a small sample of C. f. ssp. californicum was geneticall distinct (Armitage et al. 2017). More extensive study of West Coast populations is needed.

Two nominal subspecies, C. f. subsp. atlanticum, and C. f. subsp. scandinavicum coexist with C. fragile in Europe and are of uncertain taxonomic and introduction status. They are not regarded as invasive. Codium fragile subsp. atlanticum is smaller (usually 15-25 cm high) and has much smaller utricle tips than typical C. fragile (Silva 1955). Some herbarium specimens, labelled as atlanticum were genetically distinct, while others matched typical C. fragile (Provan et al. 2008). One population, morphologically resembling atlanticum, was found in Malpeque Bay, Prince Edward Island, Canada (Hubbard and Garbary 2002). Codium fragile subsp. scandinavicum has a limited reported range in Norway and Sweden. Provan et al. (2008) found that herbarium specimens of scandinavicum were genetically identical to C. fragile. Both of these subspecies may be synonymous and are best regarded as ecotypes. ‘The degree of both genotypic variation reported (small) and phenotypic variation described (large) makes the described subspecies and varieties unsustainable and it would be better to recognize one variable species for all practical purposes.’ - Wendy Guiry (29 Dec. 2007, in Guiry and Guiry 2016). A recent moleclular analysis of the DNA barcoding marker tufA suggests that there at least 2 species within the Codium fragile clade, but the authors suggest hat more genetic research is needed to divide C. fragile into native and introduced clades throughout its worldwide range (Verbruggen et al. 2016).

---

Taxonomy

Ecology

General:

Sexual reproduction in Codium fragile takes place through the production and release of flagellated swarmers (zoospores). In other species of Codium, reproduction is anisogamic and usually dioecious, with female plants producing spores and males producing smaller spores. The spores are produced by gametangia, reproductive structures protruding from the surface of the utricle. Female gametangia are dark green and lumpy, while male gametangia have an even outline and are brightly-colored (Bold and Wynne 1978). True C. fragile are monecious (Trowbridge 1998; Prince and Trowbridge 2004). Populations of C. fragile in Long Island Sound, Europe, and Japan, reproduce by stem fragmentation and by release of zoospores, of roughly equal size and believed to be female. One population in the Northwest Atlantic (Appledore Island, Maine) produces gametes (Prince 1988). Germination of the spores of all these populations occurred without the fusion of gametes and so was parthogenetic (Ramus 1971; Churchill and Moeller 1972; Bulleri et al. 2007). In Long Island Sound, the release occurs from May to September (Churchill and Moeller 1972). The spores settle and germinate about 24 hours after release and grow into masses of filaments. Mechanical shaking was necessary for growth of primordia in culture, suggesting that tides and currents are needed to stimulate growth (Ramus 1972). In Rhode Island populations, fragmentation was most common in winter, but these winter fragments are probably not viable (Hanisak 1979). In Nova Scotia populations, fragmentation occurred in summer, as a result of wave stress, and these fragments functioned as propagules (Bégin et al. 2003). Strong light exposure results in accumulation of oxygen in the thalli, and favors long-distance dispersal and colonization by the fragments (Gagnon et al. 2014).

Codium fragile is tolerant of a wide range of temperatures (-2-30°C) and salinities (12-42 PSU) (Malinowski and Ramus 1973; Hanisak 1979), but has not colonized tropical regions. It has colonized a wide range of habitats, including rocky and cobble shores, driftwood, jetties, breakwaters, seawalls, docks, piers, and floats. On soft bottoms, it grows on scattered stones and shells (Ramus 1971; Gosner 1978; Bulleri and Airoldi 2005; Geraldi et al. 2014). This seaweed has considerable tolerance to desiccation and extends into the lower intertidal zone. In experiments, it was able to survive periods of emersion of up to 90 days in high relative air humidity, leading to the likelihood on transport in fishnets, boat decks, or fouled shellfish (Schaffelke and Deane 2005). This seaweed quickly recovers normal photosynthesis after short (5-6 h) exposure to desiccation or freshwater (Kim and Garbary 2007). Codium fragile has colonized kelp beds, particularly after disturbance by storms, herbivores such as sea urchins, or the killing of kelps by the invading bryozoan Membranipora membranacea (Harris and Mathieson 2000; Chapman et al. 2002; Scheibling and Gagnon 2006). Eelgrass (Zostera marina) beds are also invaded by C. fragile, often growing on stones or shells (Ramus 1971), but also attaching to roots and rhizomes of the plant. Garbary et al. (2004) has described a growth form which attaches to eelgrass and sends out horizontal axes and vertical branches.

Growth rates of Codium fragile, at an optimum temperature, were greatest at low to moderate light intensities, 30-90 µE m-3s-1, and long photoperiods, 16h L: 8h D, or 24 h L, but fell off a bit at high intensities (150, µE m-3s-1), suggesting best growth under summer conditions. Total daily irradiance, rather than photoperiod appears to be the major feature of light that affects growth. On a daily basis, at 24 C growth rates stopped increasing above 4 µE m-3s-1 (Hanisak 1979). Sporelings were more sensitive to light, with growth saturating at 1.2 µE m-3s-1 -day and declining above 4 µE m-3s-1 -day (Hanisak 1979). Nitrate, nitrite, ammonium, and urea were equally good as nitrogen sources (Hanisak 1979).

Codium fragile is a potentially important food item for marine herbivores, a substrate for epiphytes and epifauna, and a refuge for mobile animals. For some generalist herbivores, it is a low-quality food, however. Herbivores feeding or growing poorly on this alga include the amphipod Ampithoe longimana (Cruz-Rivera and Hay 2001) and the Green Sea Urchin Strongylocentrotus droebachiensis (Prince and LeBlanc 1992; Scheibling and Anthony 2001). However, the introduced Asian Shore Crab Hemigrapsus sanguineus preferred C. fragile and Ulva spp. to Chondrus crispus, Fucus spp., and Ascophyllum nodosum (Bourdeau and O'Connor 2003). Grazing by the introduced Common Periwinkle (Littorina littorea) may control C. fragile in the intertidal zone (Scheibling et al. 2008). Saccoglossan seaslugs include specialized herbivores on green algae. Placida dendritca (probably a complex of several cryptic species) is a major herbivore on Codium fragile in the Gulf of Maine (Bleakney 1989; Trowbridge et al. 1998; Harris and Jones 2005).

Consumers:

Placida dendritica (sea slug)

Trophic Status:

Primary Producer

PrimProd

---

Habitats

General Habitat	Grass Bed	None
General Habitat	Unstructured Bottom	None
General Habitat	Oyster Reef	None
General Habitat	Marinas & Docks	None
General Habitat	Rocky	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

---

Life History

---

Tolerances and Life History Parameters

Minimum Temperature (ºC)	-2	Field data, New England (Malinowski and Ramus 1972)
Maximum Temperature (ºC)	30	Experimental data growth of thalli (Hanisak 1979)
Minimum Salinity (‰)	12	Experimental data, growth of thalli at 24 C (Hanisak 1979)
Maximum Salinity (‰)	42	Experimental data, growth of thalli at 24 C ( (Hanisak 1979)
Minimum Reproductive Temperature	12	Experimental data, growth of sporelings (Hanisak 1979)
Maximum Reproductive Temperature	24	Experimental data, growth of sporelings (Hanisak 1979)
Minimum Reproductive Salinity	18	Experimental data, growth of sporelings (Hanisak 1979)
Maximum Reproductive Salinity	48	Experimental data, growth of sporelings (Hanisak 1979)
Maximum Length (mm)	1,000	More typically to 500 mm or less (Silva 1955; Gosner 1978; Schneider and Searles 1991; Van Patten 2006)
Broad Temperature Range	None	Cold temperate-Warm temperate
Broad Salinity Range	None	Polyhaline-Euhaline

General Impacts

Codium fragile is widely regarded as a troublesome invader. The DAISIE website lists it as one of the '100 worst' invaders in Europe (DAISIE 2009), as does a species list for the Mediterranean (Streftaris and Zeneros 2006). Common economic impacts include interference with fishing gear and aquaculture, and aesthetic impacts on recreation, due to masses of the slimy weed washed up on beaches (Ramus 1971; Boudouresque 1994). Codium fragile's invasion of productive marine habitats such as kelp and seagrass beds may have impacts on fisheries, but these are difficult to measure (Ramus 1971; Schmidt and Sheibling 2006; Kelly et al. 2011).

Economic Impacts

Fisheries - Codium fragile can attach to shellfish, interfering with movement in scallops and causing shellfish, mussels, and oysters to be washed ashore with the buoyant seaweed (Ramus 1971). The weed also frequently fouls fishing lines and nets in shallow waters (Galtsoff 1964; Boudouresque 1994; Campbell 1999; Garbary et al. 2004). In Chile, it overgrew red algae (Gracilaria chilensis), being cultured for agar, reducing the harvest of the crop (Neill et al. 2006).

Aesthetic - Accumulations of Codium fragile on beaches are unattractive and unpleasant to walk on, perhaps more so than most seaweeds. On public beaches, the seaweeds are regularly removed by cleaning crews (Fofonoff personal observation). On French Mediterranean beaches, in the 1960s, blooms of C. fragile washed ashore, and had to be removed with heavy equipment (Boudouresque 1994).

Ecological Impacts

Competition - In invaded areas, C. fragile has often been reported to invade native plant communities and to partially replace native species. The extent of this competition seems to vary greatly in different regions and different plant communities. It appears to do best as a colonizer of bare surfaces, such as artificial structures, and areas cleared by storms, grazers (such as sea urchins), or the overgrowing bryozoan Membranipora membranacea, and rarely invades areas of undisturbed, established vegetation, such as kelp or seagrass beds (Malinowski and Ramus 1973; Harris and Tyrell 2001; Chapman et al. 2002; Scheibling et al. 2006). Its advantages over native seaweeds include rapid growth and longevity, since most native marine plants are annuals. It rapidly responds to added nutrients, giving it an advantage in eutrophic areas (Ramus 1971). In European waters, it was reported to be replacing the native C. tomentosum and C. vermilaria (Farnham 1980), but in a later survey, native Codium were found to dominate on natural surfaces, while C. fragile dominated on docks and breakwaters (Trowbridge and Farnham 2009). A similar pattern was seen in North Carolina, with C. fragile replacing the native C. decorticatum on artificial substrates (concrete, bulkheads, and granite seawalls), while the native species remains abundant on natural substrates such as oysters (Geraldi et al. 2014).

Habitat Change - Codium fragile is an 'ecosystem engineer', and can alter coastal habitats in multiple ways, especially in regions like the Northeast coast of North America, where no native Codium is present, and where no other alga has a similar growth form. Codium fragile, and its buoyant branches, can interfere with the movement of scallops, help waves wash mussels and stones on to beaches, and float oysters away from their beds (Ben-Avraham 1971; Ramus 1971). On the other hand, they also provide a substrate for epiphytes and epifauna, and a refuge for mobile prey, although often differing in structure, food quality, and physical characteristics from native seaweeds and sea grasses.

Codium fragile is widely thought to provide inferior habitat to native seaweeds and seagrasses, but the evidence for this is mixed. Codium fragile in the Gulf of Maine replaced kelp canopy with less favorable habitat and supported fewer juvenile fishes (Cunner, Tautogolabrus adspersus). This was attributed to the inferior food quality of C. fragile, and a scarcity of epiphytes (Levin et al. 2002). However, experiments elsewhere suggest more complex effects, with different species responding differently to changes in plant communities. In Narragansett Bay, C. fragile supported a more diverse and dense community of native and introduced epiphytic algae, than the native algae Fucus vesiculosus (Jones and Thornber 2005). In Cranberry Cove, Nova Scotia, C. fragile had a greater density (but not diversity) of epiphytes than kelps (Laminaria spp.). The differing structure of a C. fragile canopy permits greater survival of juvenile fishes and crabs than in barren regions or kelp beds (Schmidt and Scheibling 2006). However, these changes were transient in regions off Nova Scotia, where the large growths of C. fragile were removed by winter storms and recolonized by native kelps (Kelly et al. 2011). Small bivalves and amphipods were most abundant in the C. fragile-dominated state, compared to amphipods and small gastropods in the kelp-dominated state (Kelly et al. 2011). In Magdalen Islands, Quebec, eelgrass beds (Zostera marina) had greater diversity and abundance of invertebrates and fishes when invaded by C. fragile (Drouin et al. 2011). In the Adriatic Sea, C. fragile increased the recruitment and survival of Mediterranean Mussels (Mytlilus galloprovincialis) on breakwaters, offering an irregular surface, colonized by filamentous epiphytic algae, favorable for settlement, and buffering mussels from thermal stress and desiccation (Bulleri et al. 2006).

Food/Prey - For many generalized herbivores C. fragile is an inferior food. However, it can support edible epiphytic algae and it also hosts some saccoglossan slugs, which are green-alga-specialists (Trowbridge 1988). Among East Coast species showing poor feeding, growth, or reproduction on C. fragile were the snail Lacuna vincta (Northern Lacuna), the amphipod Ampithoe longimana (Cruz-Rivera and Hay 2001), and the Green Sea Urchin Strongylocentrotus droebachiensis (Prince and LeBlanc 1992; Scheibling and Anthony 2001). However, the introduced Asian Shore Crab Hemigrapsus sanguineus preferred C. fragile to several native seaweeds (Bourdeau and O'Connor 2003). The herbivorous saccoglossan sea-slug Pacifia dendritica feeds on C. fragile, and can control the seaweed's populations, but is vulnerable to predators, including the introduced Green Crab (Carcinus maenas) (Harris and Jones 2005). This cosmopolitan sea slug (probably a species complex) may have been present, but rare on the East Coast, before the Codium invasion, but is now abundant (Bleakney 1989; Harris and Jones 2005).

---

Regional Impacts

NA-ET1	Gulf of St. Lawrence to Bay of Fundy		Ecological Impact	Competition
Codium fragile invades after disturbance to kelp beds, and displaces native kelps (Laminaria digitata and L. longicruris). Major causes of displacement of kelps include outbreaks of the sea urchin Strongylocentrotus droebachiensis, and the killing of kelps by the introduced bryozoan Membranipora membranacea. A survey in the year 2000 found extensive meadows of C. fragile in Mahone and St. Margarets Bays (Chapman et al. 2002). In experiments, removal of Laminaria and a smaller native brown alga, Demarestia sp. resulted in an increase in C. fragile biomass. Conversely, removal of C. fragile resulted in increased growth of Laminaria and Demarestia species. Codium fragile meadows inhibit the recruitment of native kelps and other algae (Scheibling et al. 2006). However, in areas off the coast of Nova Scotia, large growths of C. fragile were vulnerable to winter storms, and declined from 2000 to 2008, and were replaced by the native kelp L. digitata (Kelly et al. 2011).
NA-ET1	Gulf of St. Lawrence to Bay of Fundy		Ecological Impact	Habitat Change
Codium fragile can replace a kelp canopy with less favorable habitat (Chapman et al. 2002). The differing structure of canopy of Codium fragile permits better survival of juvenile fishes and crabs (Schmidt and Sheibling 2007). These changes were transient, in regions off Nova Scotia, where the large growths of C. fragile were removed by winter storms, and recolonized by native kelps (Kelly et al. 2011). Small bivalves and amphipods were most abundant in the C. fragile-dominated state, compared to amphipods and small gastropods in the kelp-dominated state (Kelly et al. 2011).
NA-ET2	Bay of Fundy to Cape Cod		Ecological Impact	Competition
Codium fragile invades kelp beds after disturbances, and displaces kelps, and inhibits kelp recruitment. It exploits the fouling and killing of kelps by the introduced bryozoan Membranipora membranacea (Levin et al. 2002). In shallow waters along the Isle of Shoals, there was a gradual shift from kelps to Codium and to an opportunistic red alga community (Harris and Tyrell 2001).
NA-ET2	Bay of Fundy to Cape Cod		Ecological Impact	Habitat Change
Codium fragile replaces kelp canopy with less favorable habitat, and supports fewer juvenile fishes (Cunner, Tautogolabrus adspersus) (Levin et al. 2002). Around the Isles of Shoals, NH-ME, from the 1979 to 2015, there has been a trend of replacement of kelps and other larger brown seaweeds, by smaller, bushier red seaweeds (Bonnemaisonia hamifera, Dasysiphonia japonica, and Neosiphonia spp.) and Codium fragile, increasing the structural complexity of the environment, and the abundance and diversity of meso-sized invertebrates (Dijkstra et al. 2017).
NA-ET1	Gulf of St. Lawrence to Bay of Fundy		Ecological Impact	Food/Prey
Codium fragile was poor food for the sea urchin Strongylocentrotus droebachiensis. The urchins consumed 3.5X more algae than those fed Laminaria sp., but had less growth, and no gonadal production (Scheibling and Anthony 2001).
M020	Narragansett Bay		Ecological Impact	Habitat Change
Codium fragile supports a diverse community of native and introduced epiphytic algae, year-round. The latter include Antithamnion hubbsii, Bonnemaisonia hamifera, and Neosiphonia harveyi. Epiphyte communities on C. fragile were denser and more abundant than those on the native Fucus vesiculosus (Jones and Thorber 2010). The epiphytes N. harveyi (introduced) and C. virgatum (native) also provided an important food source for the native snail Lacuna vincta (Northern Lacuna) (Jones and Thornber 2005).
NA-S3	None		Ecological Impact	Habitat Change
In the Magdalen Islands, Quebec, Zostera marina seagrass beds had greater diversity and abundance of invertebrates, and greater abundance of fishes, when invaded by Codium fragile (Drouin et al. 2011).
B-II	None		Ecological Impact	Competition
In the Kattegat, C. fragile was classified as having a low level of community impacts (Zaiko et al. 2011).
M050	Great South Bay		Ecological Impact	Habitat Change
Survival of young Bay Scallops (Argopecten irradians) did not differ greatly between Eelgrass (Zostera marina beds and patches of Codium fragile in Sag Harbor, Long Island. A longer term study in Sag Harbor and Shinnecock Inlet found some evidence for impacts for differences in growth rates and condition between scallop populations in eelgrass and Codium beds, but this appeared to vary among bays and years (Carroll and Peterson 2013).
NA-ET3	Cape Cod to Cape Hatteras		Ecological Impact	Habitat Change
Research in the 1970s suggested that Codium fragile may have the potential to change a sandy beach to a gravel beach (Nobska Beach, Woods Hole) by attaching and transporting stones onto shore (with it's buoyant fronds) (Ben-Avraham 1971). However this beach was still mostly sand in 2015 (Paul Fofonoff, personal observation). Codium fragile is believed to have affected the growth and mortality of Bay Scallops (Argopecten irradians), Blue Mussels (Mytilus edulis), and Eastern Oysters (Crassostrea virginica). Attached C. fragile can inhibit the movement of scallops, by attaching to their shells, causing them to float in the water column and possibly be washed ashore by waves (Ramus 1971). Survival of young Bay Scallops (Argopecten irradians) did not differ greatly between Eelgrass (Zostera marina) beds and patches of Codium fragile in Sag Harbor, Long Island. A longer term study in Sag Harbor and Shinnecock Inlet found some evidence for impacts for differences in growth rates and condition between scallop populations in eelgrass and Codium beds, but this appeared to vary among bays and years (Carroll and Peterson 2013). Codium fragile supports a diverse community of native and introduced epiphytic algae, year-round. The latter include Antithamnion hubbsii, Bonnemaisonia hamifera, and Neosiphonia harveyi. Epiphyte communities on C. fragile were denser and more abundant than those on the native Fucus vesiculosus (Jones and Thorber 2010). The epiphytes N. harveyi(introduced) and C. virgatum (native) also provided an important food source for the native snail Lacuna vincta (Northern Lacuna) (Jones and Thornber 2005).
S030	Bogue Sound		Ecological Impact	Habitat Change
Codium spp. are associated with both nitrogen-fixing and denitrifying bacteria. Codium fragile, which prefers artificial substrates, supports a greater rate of denitrification than the native C. decorticatum, increasing N2 release, and decreasing nitrate concentrations, potentially decreasing eutrophication (Geraldi et al. 2014).
S050	Cape Fear River		Ecological Impact	Habitat Change
Codium spp. are associated with both nitrogen-fixing and denitrifying bacteria. Codium fragile, which prefers artificial substrates, supports a greater rate of denitrification than the native C. decorticatum, increasing N2 release, and decreasing nitrate concentrations, potentially decreasing eutrophication (Geraldi et al. 2014). In experiments, C. fragile in single-species treatments supported a lower diversity of consumers, possibly because of lower habitat complexity, or because of lower food quality (Ramus and Long 2016).
CAR-VII	Cape Hatteras to Mid-East Florida		Ecological Impact	Habitat Change
Codium spp. are associated with both nitrogen-fixing and denitrifying bacteria. Codium fragile, which prefers artificial substrates, supports a greater rate of denitrification than the native C. decorticatum, increasing N2 release, and decreasing nitrate concentrations, potentially decreasing eutrophication (Geraldi et al. 2014). In experiments, C. fragile in single-speacies treatments supported a lower diversity of consumers, possibly because of lower habitat complexity, or because of lower food quality (Ramus and Long 2016).
AR-V	None		Ecological Impact	Competition
Codium fragile competes for space with the native brown seaweed Fucus serratus, but only in the lower subtidal part of F. serratus' range. Codium fragile tends to replace F. serratus in sheltered subtidal environments, and stony substrate. Fucus serratus is more successful on bedrock, in exposed sites (Armitage and Sjotun 2014).
AUS-VIII	None		Economic Impact	Fisheries
Codium fragile was discovered in Corner Inlet, Victoria, when it was fouling fishing nets (Campbell 1999).
S030	Bogue Sound		Ecological Impact	Food/Prey
Codium fragile was found to be a poor food source for the amphipod Ampithoe longimana, resulting in lower growth and survivorship (Cruz-Ribera and Hay 2001).
CAR-VII	Cape Hatteras to Mid-East Florida		Ecological Impact	Food/Prey
Codium fragile was found to be a poor food source for the amphipod Ampithoe longimana, resulting in lower growth and survivorship (Cruz-Ribera and Hay 2001).
S050	Cape Fear River		Ecological Impact	Competition
Codium fragile appears to have replaced the native C. decorticatum on artificial substrates (concrete, bulkheads, granite seawalls) in Cape Fear and Masonboro Sounds, in comparisons with 1940s surveys. However, C. decorticatum remains abundant on natural substrates such as oysters (Geraldi et al. 2014).
S045	_CDA_S045 (New)		Ecological Impact	Competition
Codium fragile appears to have replaced the native C. decorticatum on artificial substrates (concrete, bulkheads, granite seawalls) in Cape Fear and Masonboro Sounds, in comparisons with 1940s surveys. However, C. decorticatum remains abundant on natural substrates such as oysters (Geraldi et al. 2014).
S045	_CDA_S045 (New)		Ecological Impact	Habitat Change
Codium spp. are associated with both nitrogen-fixing and denitrifying bacteria. Codium fragile, which prefers artificial substrates, supports a greater rate of denitrification than the native C. decorticatum, increasing N2 release, and decreasing nitrate concentrations, potentially decreasing eutrophication (Geraldi et al. 2014).
CAR-VII	Cape Hatteras to Mid-East Florida		Ecological Impact	Competition
Codium fragile appears to have replaced the native C. decorticatum on artificial substrates (concrete, bulkheads, granite seawalls) in Cape Fear and Masonboro Sounds, in comparisons with 1940s surveys. However, C. decorticatum remains abundant on natural substrates such as oysters (Geraldi et al. 2014).
NEA-II	None		Ecological Impact	Competition
Codium was reported to replace the native C. tomentosum on the south and west coasts of the British Isles (Farnham 1980; Eno et al. 1997). However, in the English Channel region, dense populations of C. fragile are seen on artificial structures, such as docks and breakwaters, but populations of the native C. tomentosum and C. vermilara persist in natural habitats (Trowbridge and Farnham 2009).
NA-S3	None		Economic Impact	Fisheries
Codium fragile fouled about 80% of the space on mussel aquaculture lines (Garbary et al. 2004).
N135	_CDA_N135 (Piscataqua-Salmon Falls)		Ecological Impact	Competition
Codium fragile invades kelp beds after disturbances, and displaces kelps and inhibits kelp recruitment. It exploits the fouling and killing of kelps by the introduced bryozoan Membranipora membranacea (Levin et al. 2002). In shallow waters along the Isle of Shoals, there was a gradual shift from kelps to Codium and to an opportunistic red alga community (Harris and Tyrell 2001).
N135	_CDA_N135 (Piscataqua-Salmon Falls)		Ecological Impact	Habitat Change
Codium fragile replaces kelp canopy with less favorable habitat, and supports fewer juvenile fishes (Cunner, Tautogolabrus adspersus) (Levin et al. 2002). Around the Isles of Shoals, NH-ME, from the 1979 to 2015, there has been a trend of replacement of kelps and other larger brown seaweeds, by smaller, bushier red seaweeds (Bonnemaisonia hamifera, Dasysiphonia japonica, and Neosiphonia spp.) and Codium fragile, increasing the structural complexity of the environment, and the abundance and diversity of meso-sized invertebrates (Dijkstra et al. 2017).
N135	_CDA_N135 (Piscataqua-Salmon Falls)		Ecological Impact	Food/Prey
The native snail Lacuna vincta occurred in lower density on Codium fragile than on the native kelp Laminaria saccharina. In experiments, feeding and growth was reduced in L. vincta on Codium, compared to Laminaria (Chavanich and Harris 2004). Codium fragile was a less-preferred food of the Green Sea Urchin Strongylocentrotus droebachiensis (Prince and Leblanc 1993) The herbivorous saccoglossan sea-slug feeds on C. fragile, and can control the seaweed's populations (Harris and Adams 2005).
NA-ET2	Bay of Fundy to Cape Cod		Ecological Impact	Food/Prey
The native snail Lacuna vincta occurred in lower density on Codium fragile than on the native kelp Laminaria saccharina. In experiments, feeding and growth was reduced in L. vincta on Codium, compared to Laminaria (Chavanich and Harris 2004). The herbivorous saccoglossan sea-slug Pacifia dendritica feeds on C. fragile, and can control the seaweed's populations (Harris and Adams 2005). This sea slug may have been present, but rare on the East Coast, before the Codium invasion (Bleakney 1989).
N195	_CDA_N195 (Cape Cod)		Ecological Impact	Habitat Change
Codium fragile attaches to stones, and was observed to increase the transport of stones onto a sand beach, with the potential to change it to a gravel beach (Nobska Beach, Woods Hole) (Ben-Avraham 1971). However this beach was still mostly sand in 2015 (Paul Fofonoff, personal observation).
M023	_CDA_M023 (Narragansett)		Ecological Impact	Habitat Change
Codium fragile attached to Blue Mussels (Mytilus edulis caused large quantities of mussels to be washed onto the Beach at Charlestown, Rhode Island (Ramus 1971).
M040	Long Island Sound		Ecological Impact	Habitat Change
Codium fragile attached to the shells of Bay Scallops (Argopecten irradians) in the Niantic River, Connecticut, interfered with the movement of the scallops, and increased their mortality (Ramus 1971).
N185	_CDA_N185 (Cape Cod)		Ecological Impact	Habitat Change
Codium fragile attached to Eastern Oysters (Crassostrea virignica) caused oysters to be floated away from their beds, when the algae became saturated with oxygen, and lifted the oysters off the bottom (Ramus 1971).
N185	_CDA_N185 (Cape Cod)		Economic Impact	Fisheries
Mortality caused by Codium fragile is believed to have raised the priced of local oysters on Cape Cod. Codium fragile also interfered with oyster-harvesting equipment (Galtsoff 1965; Ramus 1971).
M040	Long Island Sound		Economic Impact	Fisheries
Codium fragile is believed to have contributed to the mortality of Bay Scallops in Long Island Sound. However, the role of C. fragile is difficult to determine, since the seaweed also attaches to dead shells (Ramus 1971).
NA-ET3	Cape Cod to Cape Hatteras		Economic Impact	Fisheries
Codium fragile is believed to have contributed to the mortality of Bay Scallops in Long Island Sound. However, the role of C. fragile is difficult to determine, since the seaweed also attaches to dead shells (Ramus 1971). Mortality caused by Codium fragile is believed to have raised the priced of local oysters on Cape Cod. Codium fragile also interfered with oyster-harvesting equipment (Galtsoff 1965; Ramus 1971).
MED-II	None		Economic Impact	Fisheries
Codium fragile has fouled ropes used in oyster culture in the Thau Lagoon (Boudouresque 1994).
MED-VII	None		Ecological Impact	Habitat Change
The juvenile thalli (primordia) and the adult canopies of Codium fragile both had positive effects on the recruitment and survival of Mediterranean Mussels (Mytlilus galloprovincialis) on breakwaters in the Adriatic Sea. The primordia offered an irregular surface, and may have altered flow, and encouraged filamentous algae, favoring settlement of larvae. The mature canopy may also buffer mussels from thermal stress and desiccation (Bulleri et al. 2006).
N195	_CDA_N195 (Cape Cod)		Economic Impact	Aesthetic
Accumulations of Codium fragile on Cape Cod beaches are unattractive and unpleasant to walk on, perhaps more so than most seaweeds (Fofonoff personal observation).
NA-ET3	Cape Cod to Cape Hatteras		Economic Impact	Aesthetic
Accumulations of Codium fragile on Cape Cod beaches are unattractive and unpleasant to walk on, perhaps more so than most seaweeds (Fofonoff personal observation).
NA-S3	None		Ecological Impact	Competition
In Caribou Harbour, Nova Scotia, a distinct horizontal growth form of Codium fragile was attaching to rhizomes of Eelgrass, potentially competing with the grass (Garbary et al. 2004). However, in the Magdalen Islands, experiments indicated that negative effects on eelgrass occurred only at high Codium densities (Drouin et al. 2013). Disturbance and exposure, of Eelgrass rhizomes by storms, ice scour, and boat traffic, appears to be a major factor controlling invasion of the grass beds by C. fragile (Gagnon et al. 2014).
SEP-C	None		Economic Impact	Fisheries
Codium fragile interferes with the aquaculture of the native red alga Gracilaria chilensis, significantly reducing the harvest of the crop (Neill et al. 2006).
NA-ET3	Cape Cod to Cape Hatteras		Ecological Impact	Food/Prey
The Asian Shore Crab Hemigrapsus sanguineus preferred C. fragile and Ulva spp. to Chondrus crispus, Fucus spp., and Ascophyllum nodossum (Bourdeau and O'Connor 2003).
M010	Buzzards Bay		Ecological Impact	Food/Prey
The Asian Shore Crab Hemigrapsus sanguineus preferred C. fragile and Ulva spp. to Chondrus crispus, Fucus spp., and Ascophyllum nodossum (Bourdeau and O'Connor 2003).
MED-II	None		Economic Impact	Aesthetic
In the 1960s, heavy depositions of Codium fragile on the shore near Marseille required removal with heavy equipment (Boudouresque 1994).
M128	_CDA_M128 (Eastern Lower Delmarva)		Ecological Impact	Competition
In Hog Island Bay, Virginia, Codium fragile was competitive on hard substrates where sediment and drift algae did not accumulate, with moderate growth rates, but a long growing season. It was the fourth most abundant alga in Hog Island Bay (Thomsen 2003).
NA-ET3	Cape Cod to Cape Hatteras		Ecological Impact	Competition
In Long Island Sound, Codium fragile cannot invade established Eelgrass (Zostera marina) or native algal communities, but if an area is disturbed and cleared  C. fragile can establish (Malinowski and Ramus 1971). In Hog Island Bay, Virginia, C. fragile was competitive on hard substrates where sediment and drift algae did not accumulate, and had moderate growth rates but a long growing season. It was the fourth most abundant algae in Hog Island Bay (Thomsen 2003).
AR-V	None		Ecological Impact	Habitat Change
Epiphytic and epifaunal communities of the introduced C. fragile, off Norway, shared many species with the native Fucus serratus but showed statistical differences in composition. Fucus serratus support more sessile fauna, especially bryozoans (Armitage and Sjotun 2016).
AUS-X	None		Ecological Impact	Habitat Change
The non-native subspecies of Codium fragile supports a lower diversity but a higher abundance of amphipods (mostly Hyale spp. and the native gastropod Alaba opiniosa. Abundance and diversity of amphipods were affected by the number of branches of the seaweeds, as much as by species identity (Lutz et al. 2019).
NH	New Hampshire		Ecological Impact	Competition
Codium fragile invades kelp beds after disturbances, and displaces kelps and inhibits kelp recruitment. It exploits the fouling and killing of kelps by the introduced bryozoan Membranipora membranacea (Levin et al. 2002). In shallow waters along the Isle of Shoals, there was a gradual shift from kelps to Codium and to an opportunistic red alga community (Harris and Tyrell 2001).
NH	New Hampshire		Ecological Impact	Food/Prey
The native snail Lacuna vincta occurred in lower density on Codium fragile than on the native kelp Laminaria saccharina. In experiments, feeding and growth was reduced in L. vincta on Codium, compared to Laminaria (Chavanich and Harris 2004). Codium fragile was a less-preferred food of the Green Sea Urchin Strongylocentrotus droebachiensis (Prince and Leblanc 1993) The herbivorous saccoglossan sea-slug feeds on C. fragile, and can control the seaweed's populations (Harris and Adams 2005).
NH	New Hampshire		Ecological Impact	Habitat Change
Codium fragile replaces kelp canopy with less favorable habitat, and supports fewer juvenile fishes (Cunner, Tautogolabrus adspersus) (Levin et al. 2002). Around the Isles of Shoals, NH-ME, from the 1979 to 2015, there has been a trend of replacement of kelps and other larger brown seaweeds, by smaller, bushier red seaweeds (Bonnemaisonia hamifera, Dasysiphonia japonica, and Neosiphonia spp.) and Codium fragile, increasing the structural complexity of the environment, and the abundance and diversity of meso-sized invertebrates (Dijkstra et al. 2017).
NY	New York		Ecological Impact	Habitat Change
Survival of young Bay Scallops (Argopecten irradians) did not differ greatly between Eelgrass (Zostera marina beds and patches of Codium fragile in Sag Harbor, Long Island. A longer term study in Sag Harbor and Shinnecock Inlet found some evidence for impacts for differences in growth rates and condition between scallop populations in eelgrass and Codium beds, but this appeared to vary among bays and years (Carroll and Peterson 2013).
MA	Massachusetts		Ecological Impact	Food/Prey
The Asian Shore Crab Hemigrapsus sanguineus preferred C. fragile and Ulva spp. to Chondrus crispus, Fucus spp., and Ascophyllum nodossum (Bourdeau and O'Connor 2003).
MA	Massachusetts		Ecological Impact	Habitat Change
Codium fragile attached to Eastern Oysters (Crassostrea virignica) caused oysters to be floated away from their beds, when the algae became saturated with oxygen, and lifted the oysters off the bottom (Ramus 1971)., Codium fragile attaches to stones, and was observed to increase the transport of stones onto a sand beach, with the potential to change it to a gravel beach (Nobska Beach, Woods Hole) (Ben-Avraham 1971). However this beach was still mostly sand in 2015 (Paul Fofonoff, personal observation).
MA	Massachusetts		Economic Impact	Aesthetic
Accumulations of Codium fragile on Cape Cod beaches are unattractive and unpleasant to walk on, perhaps more so than most seaweeds (Fofonoff personal observation).
MA	Massachusetts		Economic Impact	Fisheries
Mortality caused by Codium fragile is believed to have raised the priced of local oysters on Cape Cod. Codium fragile also interfered with oyster-harvesting equipment (Galtsoff 1965; Ramus 1971).
RI	Rhode Island		Ecological Impact	Habitat Change
Codium fragile attached to Blue Mussels (Mytilus edulis caused large quantities of mussels to be washed onto the Beach at Charlestown, Rhode Island (Ramus 1971).
VA	Virginia		Ecological Impact	Competition
In Hog Island Bay, Virginia, Codium fragile was competitive on hard substrates where sediment and drift algae did not accumulate, with moderate growth rates, but a long growing season. It was the fourth most abundant alga in Hog Island Bay (Thomsen 2003).
NC	North Carolina		Ecological Impact	Competition
Codium fragile appears to have replaced the native C. decorticatum on artificial substrates (concrete, bulkheads, granite seawalls) in Cape Fear and Masonboro Sounds, in comparisons with 1940s surveys. However, C. decorticatum remains abundant on natural substrates such as oysters (Geraldi et al. 2014)., Codium fragile appears to have replaced the native C. decorticatum on artificial substrates (concrete, bulkheads, granite seawalls) in Cape Fear and Masonboro Sounds, in comparisons with 1940s surveys. However, C. decorticatum remains abundant on natural substrates such as oysters (Geraldi et al. 2014).
NC	North Carolina		Ecological Impact	Food/Prey
Codium fragile was found to be a poor food source for the amphipod Ampithoe longimana, resulting in lower growth and survivorship (Cruz-Ribera and Hay 2001).
NC	North Carolina		Ecological Impact	Habitat Change
Codium spp. are associated with both nitrogen-fixing and denitrifying bacteria. Codium fragile, which prefers artificial substrates, supports a greater rate of denitrification than the native C. decorticatum, increasing N2 release, and decreasing nitrate concentrations, potentially decreasing eutrophication (Geraldi et al. 2014)., Codium spp. are associated with both nitrogen-fixing and denitrifying bacteria. Codium fragile, which prefers artificial substrates, supports a greater rate of denitrification than the native C. decorticatum, increasing N2 release, and decreasing nitrate concentrations, potentially decreasing eutrophication (Geraldi et al. 2014)., Codium spp. are associated with both nitrogen-fixing and denitrifying bacteria. Codium fragile, which prefers artificial substrates, supports a greater rate of denitrification than the native C. decorticatum, increasing N2 release, and decreasing nitrate concentrations, potentially decreasing eutrophication (Geraldi et al. 2014). In experiments, C. fragile in single-species treatments supported a lower diversity of consumers, possibly because of lower habitat complexity, or because of lower food quality (Ramus and Long 2016).
ME	Maine		Economic Impact	Fisheries
Clogging fishnets and lobsterpots (Fisherman's comment at a meeting, heard by Paul Fofonoff)

References

´Bourmaud, Chloe; Gravier-Bonnet, Nicole (2004) Medusoid release and spawning of Macrorynchia philippina Kirchenpauer, 1872 (Cnidaria, Hydrozoa, Aglaopheniidae), Hydrobiologia 530/531: 365–372,

Abbott, Isabella A.; Hollenberg, George J. (1976) <missing title>, Stanford University Press, Stanford CA. Pp. <missing location>

Adey, Walter H.; Hayek, Lee-Ann C. (2011) Elucidating marine biogeography with macrophytes: Quantitative analysis of the North Atlantic supports the thermogeographic model and demonstrates a distinct Subarctic region in the Northwestern Atlantic, Northeastern Naturalist 18(8): 1-128

Aguirre-Pabon. Juan; Chasqui, Luis; Munoz, Eider; Narvaez-Barandica, Juan (2023) Multiple origins define the genetic structure of tiger shrimp Penaeus monodon in the colombian Caribbean Sea, Heliyon 9(e17727): Published online
https://doi.org/10.1016/j.heliyon.2023.e17727

Airoldi, Laura; Bulleri, Fabio (2011) Anthropogenic disturbance can determine the magnitude of opportunistic species responses on marine urban infrastructures, PLOS ONE 6(8): e22985

Appeltans, W. et al. 2011-2015 World Registry of Marine Species. <missing URL>



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

Armitage, Caroline S.; Sjøtun, Kjersti; Jensen, Knut Helge (2014) Correlative evidence for competition between Fucus serratus and the introduced chlorophyte Codium fragile subsp. fragile on the southwest coast of Norway, Botanica Marina 57(2): 85-97

Ashton, Gail ; Boos, Karin; Shucksmith, Richard; Cook, Elizabeth (2006) Rapid assessment of the distribution of marine non-native species in marinas in Scotland, Aquatic Invasions 1(4): 209-213

Associated Press (12/2021) Lummi Nation declares disaster after invasive crab arrives, Seattle Times <missing volume>: <missing location>

Baldwin, Andy; Leason, Diane (2016) Potential Ecological impacts of Emerald Ash Borer on Maryland's Eastern Shore, In: None(Eds.) None. , <missing place>. Pp. <missing location>

Banagouro, Quaglio; Karine Cristie; Viana, Jefferson; Pereira de Lima, Leonardo; Coelho, Rocha, Thalita; Girardello, Guilherme Rabelo; Russi, Karolayne Larissa; Kitahara , Marcelo V.; Marcelo V. Sciani, Juliana Mozer ; (2021) Biochemical and Toxinological Characterization of Venom from Macrorhynchia philippina (Cnidaria, Hydrozoa), BioMed Research 2022: Published online
https://doi.org/10.1155/2022/8170252

Barbara, I.; Cremades, J. (1996) Seaweeds of the Ria de A Coruna (NW Iberian Peninsula, Spain), Botanica Marina 39: 371-388

Bazairi, H. and 7 authors (2013) Alien marine species of Libya: first inventory and new records in El-Kouf National Park (Cyrenaica) and the neighbouring areas, Mediterranean Marine Science 14(2): 451-462

Bégin, Chantale; Scheibling, Robert E. (2003) Growth and survival of the invasive green alga Codium fragile ssp. tomentosoides in tide pools on a rocky shore in Nova Scotia, Botanica Marina 46: 404-412

Ben-Avraham, Zvi (1971) Accumulation of stones on beaches by Codium fragilie, Limnology and Oceanography 16: 553-554

Bernardi, Giacomo; Cohn, Francesca; Dominguez‑Dominguez, Omar; Kingon, Kelly; Tornabene, Luke; Robertson, D. Ross (2024) Establishment genomics of the Indo‑Pacific damselfish Neopomacentrus cyanomos, in the Greater Caribbean, Biological Invasions <missing volume>: Published online
https://doi.org/10.1007/s10530-023-03226-w

Bird, C. J., Dadswell, M. J., Grund, D. W. (1993) First record of the potential nuisance alga Codium fragile ssp. tomentosoides (Chlorophyta, Caulerpales) in Atlantic Canada, Proceedings of the Nova Scotian Institute of Science 40: 11-17

Bird, C. J.; Dadswell, M. J.; Grund, D. W. (1993) First record of the potential nuisance alga Codium fragile spp. tomentosoides (Chlorophyta, Caulerpales) in Atlantic Canada, Proceedings of the Nova Scotian Institute of Science 40: 11-17

Blanco, Andreu; Beger, Maria; Olabarria, Celia (2021) First confirmed occurrence of Codium fragile (Suringar) Hariot in the Iberian Peninsula coast of Portugal, BioInvasions Records 10: <missing location>

Bleakney, J. Sherman (1989) Morphological variation in the radula of Placida dendritica (Alder & Hancock, 1843) (Opisthobranchia: Ascoglossa/Saccoglossa) from Atlantic and Pacific populations, The Veliger 32(2): 171-181

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

Booth, David; Provan, Jim, Maggs, Christine A. (2007) Molecular approaches to the study of invasive seaweeds., Botanica Marina 50: 385396

Bouck, G. B.; Morgan, E. (1957) The occurrence of Codium in Long Island waters, Bulletin of the Torrey Botanical Club 84(5): 384-387

Boudouresque, C. F., Briand, F., Nolan, C. (1994) <missing title>, European Commission, Brussels. Pp. <missing location>

Boudouresque, Charles F. (1994) Les espèces introduites dans les eaux côtières d'Europe et de Méditerranée: Etat de la question et conséquences, In: Boudouresque, C. F.//Briand, F.//Nolan, C.(Eds.) Introduced Species in European Coastal Waters. , Brussels. Pp. 8-27

Bourdeau, Paul E.; O'Connor, Nancy J. (2003) Predation by the nonindigenous Asian Shore Crab Hemigrapsus sanguineus on macroalgae and molluscs., Northeastern Naturalist 10(3): 319-334

Bulleri, F.; Airoldi, L.; Branca, G.M.; Abbiati, M. (2005) Positive effects of the introduced green alga, Codium fragile ssp. tomentosoides, on recruitment and survival of mussels., Marine Biology 148: 1213-1220

Bulleri, Fabio; Airoldi, Laura (2005) Artificial marine structures facilitate the spread of a non-indigenous green alga, Codium fragile ssp. tomentosoides in the north Adriatic Sea., Journal of Applied Ecology 42: 1063-1072

Bulleri, Fabio; Branca, Maria Giovanna; Abbiati, Marco; Airoldi, Laura (2007) Development of reproductive structures in the introduced green alga, Codium fragile ssp. tomentosoides, in the northern Adriatic Sea, European Journal of Phycology 42(2): 37-144

Buschbaum, Christian; Lackschewitz, Dagmar; Reise, Karsten (2012) Nonnative macrobenthos in the Wadden Sea ecosystem, Journal of Ocean Management 68: 89-101

Campbell, Stuart J. (1999) Occurrence of Codium fragile subsp. tomentosoides (Chloropqhyta: Bryopsidales) in marine embayments of southeastern Australia, Journal of Phycology 35: 938-940

Cardigos, F. and 5 authors (2006) Non-indigenous marine species of the Azores., Helgoland Journal of Marine Research 60: 160-169

Carlton, J.T.; Scanlon, J.A. (1985) Progression and dispersal of an introduced alga: Codium fragile ssp. tomentosoides (Chlorophyta) on the Atlantic Coast of North America., Botanica Marina 28: 155-165

Carroll, John M.; Peterson, Bradley J.; Bonal, Dennis; Weinstock, Andrew; Smith, Christopher F.; Tettelbach, Stephen T. (2010) Comparative survival of bay scallops in eelgrass and the introduced alga, Codium fragile, in a New York estuary, Marine Biology 157: 249-259

Carroll, John M.; Peterson, Bradley J. (2013) Comparisons in demographic rates of bay scallops in eelgrass and the introduced alga, Codium fragile, in New York, Marine Biology 160: 1451-1463

Castilla, Juan C. and 10 authors (2005) Down under the southeastern Pacific: marine non-indigenous species in Chile., Biological Invasions 7: 213-232

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>

Chapman, Annelise S.; Scheibling, Robert E. Chapman, Anthony R. O. (2002) Alien invaders: In Canada's waters, wetlands, and forests, Natural Resources Canada, Ottawa, Ontario. Pp. 133-148

Chapman, Annelise Sabine (1999) From introduced species to invader: what determines variation in the success of Codium fragile ssp. tomentosoides (Chlorophyta) in the North Atlantic Ocean?, Helgoländer Meeresuntersuchungen 52: 277-289

Chavanich, Suchana; Harris, Larry G.; Je, Jong-Geel; Kang, Rae-Seon (2006) Distribution pattern of the green alga Codium fragile (Suringar) Hariot, 1889 in its native range, Korea., Aquatic Invasions 1(3): 99-108

Chavanich, Suchana; Harris, Larry G. (2004) Impact of the non-native macroalga Codium fragile (Sur.) ssp. tomentosoides (van Goor) Silva on the native snail Lacuna vincta (Montagu, 1803) in the Gulf of Maine., Veliger 47(2): 85-90

Churchill, A. Coolidge; Moeller, Henry W. (1972) Seasonal patterns of reproduction in New York populations of Codium fragile (Sur.) subsp. tomentosoides (Van Goor) Silva, Journal of Phycology 8: 147-152

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

Cohen, Andrew N. and 10 authors (2005) <missing title>, San Francisco Estuary Institute, Oakland CA. Pp. <missing location>

Cohen, Andrew N.; Carlton, James T. (1995) Nonindigenous aquatic species in a United States estuary: a case study of the biological invasions of the San Francisco Bay and Delta, U.S. Fish and Wildlife Service and National Sea Grant College Program (Connecticut Sea Grant), Washington DC, Silver Spring MD.. Pp. <missing location>

Coleman, Annette W. (1996) DNA analysis methods for recognizing species invasion: the example of Codium, and generally applicable methods for algae, Hydrobiologia <missing volume>: 29-34

Collin, Samuel B.; Tweddle, Jacqueline F.; Shucksmith, Rachel J. (2015) Rapid assessment of marine non-native species in the Shetland Islands, Scotland, BioInvasions Records 4: In press

Cordone, Georgina; Galván, David E.; Momo, Fernando R. (2023) Impacts of an invasion by green crab Carcinus maenas on the intertidal food web of a Patagonian rocky shore, Argentina, Marine Ecology Progress Series 713: 97-115
https://doi.org/10.3354/meps14336

Cranfield, H.J.; Gordon, D.P.; Willan, R.C.; Marshall, B.A; Battershill, C.N.; Francis, M.P.; Nelson, W.A.; Glasby, C.J.; Read, G.B. (1998) <missing title>, The National Institute of Water and Atmospheric Research, New Zealand. Pp. <missing location>

Cruz-Rivera; Hay, Mark E. (2001) Macroalgal traits and the feeding and fitness of an herbivorous amphipod: the roles of selectivity, mixing, and compensation, Marine Ecology Progress Series 218: 249-266

DAISIE (Delivering Alien Invasive Species Inventories to Europe) (2009) Handbook of alien species in Europe, Springer, Dordrecht, Netherlands. Pp. 269-374

Davidson, Alisha D.; Campbell, Marnie L.; Hewitt, Chad L.; Schaffelke, Britta (2015) Assessing the impacts of nonindigenous marine macroalgae: an update of current knowledge, Botanica Marina 58(2): 55-79

Di Martino, Vincenzo; Stancanelli, Bessy (2021) The alien lionfish, Pterois miles (Bennett, 1828), enters the Adriatic Sea, Central Mediterranean Sea, Journal of the Black Sea/Mediterranean Environment 27(1): 104-108

Dromgoole, F. I., Foster, B. A. (1983) Changes to the marine biota of the Auckland Harbour, Tane 29: 79-96

Drouin, Annick; McKindsey, Christopher W.; Johnson, Ladd E. (2012) Detecting the impacts of notorious invaders: experiments versus observations in the invasion of eelgrass meadows by the green seaweed Codium fragile, Oecologia 168: 491-502

Drouin, Annick; McKindsey, Christopher W.; Johnson, Ladd E. (2011) Higher abundance and diversity in faunal assemblages with the invasion of Codium fragile ssp. fragile in eelgrass meadows, Marine Ecology Progress Series 424: 105-117

Eno, N. Clare (1996) Non-native marine species in British waters: effects and controls, Aquatic Conservation: Marine and Freshwater Ecosystems 6: 215-228

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

Eno, N.C., Clark, R.A., Sanderson, W.G. 1997-2012 Directory of Non-Native Marine Species in British waters. <missing URL>



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

Fisheries and Oceans Canada 2011 Aquatic invasive species identification Booklet. <missing URL>



Fralick, R. A., Mathieson, A. C. (1973) Ecological studies of Codium fragile in New England, USA, Marine Biology 19: 127-132

Gagnon, Karine; McKindsey, Christopher W.; Johnson, Ladd E. (2014) Roles of dispersal mode, recipient environment and disturbance in the secondary spread of the invasive seaweed Codium fragile, Biological Invasions Published online: <missing location>

Gagnon, Karine; McKindsey, Christopher W.; Johnson, Ladd E. (2011) Dispersal potential of invasive algae: the determinants of buoyancy in Codium fragile ssp. fragile, Marine Biology <missing volume>: published online

Gagnon, Karine; Peacock, Stephanie J.; Jin, Yu; Lewis, Mark A. (2015) Modelling the spread of the invasive alga Codium fragile driven by long-distance dispersal of buoyant propagules, Ecological Modelling 316: 111-121

Galtsoff, Paul S. (1964) The American oyster, Crassostrea virginica Gmelin. Chapter 18: Environmental factors affecting oyster populations, Fisheries Bulletin 64: 397-456

Garbary, D. J.; Vandermeulen, H.; Kim, K. Y. (1997) Codium fragile spp. tomentosoides (Chlorophyta) invades the Gulf of Lawrence, Atlantic Canada., Botanica Marina 40: 539-540

Garbary, David J.; Fraser, Sarah J.; Hubbard, Charlene.; Kim, Kwang Young (2004) Codium fragile: rhizomatous growth in the Zostera thief of eastern Canada, Helgoland Marine Research 58: 141-146

Garbary, David J.; Miller, Anthony G.; Williams, Jim; Seymour, Norm R. (2014) Drastic decline of an extensive eelgrass bed in Nova Scotia due to the activity of the invasive green crab (Carcinus maenas), Marine Biology 161: 3-15

Geraldi, Nathan R.; Smyth, Ashley R.; Piehler, Michael F.; Peterson, Charles H. (2014) Artificial substrates enhance non-native macroalga and N2 production, Biological Invasions 16: 1819-1831

Godwin, L. Scott 1997 conversation with Paul Fofonoff. <missing URL>



Goff, Lynda J.; Liddle, Larry; Silva, Paul C.; Voytek, Mary; Coleman, Annette (1992) Tracing species invasion in Codium, a siphonous green alga, using molecular tools, American Journal of Botany 79(11): 1279-1285

Golikov, A. N. and 7 other editors. (1976) <missing title>, Nauk, Leningrad. Pp. <missing location>

Gosner, Kenneth L. (1978) A field guide to the Atlantic seashore., In: (Eds.) . , Boston. Pp. <missing location>

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

GRAJALES, ALEJANDRO; RODRÍGUEZ, ESTEFANÍA (2014) Morphological revision of the genus Aiptasia and the family Aiptasiidae (Cnidaria, Actiniaria, Metridioidea), Zootaxa 3826(1): <missing location>
https://doi.org/10.11646/zootaxa.3826.1.2

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

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



Hanisak, M. D. (1979) Growth patterns of Codium fragile tomentosoides in response to temperature, irradiance, salinity, and nitrogen source, Marine Biology 50: 319-332

Haroun, R. J.; Gil-RodrÌguez, M. C.; de Castro, J. DÌaz; Prud'homme van Reine, W. F. (2002) A checklist of the marine plants from the Canary Islands (Central Eastern Atlantic Ocean), Botanica Marina 45: 139-169

Harris, Courtney E.; K. Vogelbein, Wolfgang (2006) Parasites of Mummichogs, Fundulus heteroclitus, from the York River, Virginia, U.S.A., with a Checklist of Parasites of Atlantic Coast Fundulus spp., Comparative Parasitology 73(1): 72-110
https://doi.org/10.1654/41716.1

Harris, Larry G.; Jones, Adam C. (2005) Temperature, herbivory and epibiont acquisition as factors controlling the distribution and ecological role of an invasive seaweed., Biological Invasions 7: 913-924.

Harris, Larry G.; Mathieson, Arthur C. (2000) Patterns of range expansion, niche shift and predator acquisition in the Gulf of Maine, In: Pederson, Judith(Eds.) Marine Bioinvasions. , Cambridge. Pp. 46-56

Harris, Larry; Tyrrell, Megan (2001) Changing community states in the Gulf of Maine., Biological Invasions 3: 9-21

Hastings, Anna B. (1930) Cheilostomatous Polyzoa from the vicinity of the Panama Canal, collected by Dr, C. Crossland on the cruise of S. Y. St. george, Proceedings of the Zoological Society of London 47: 697–750

Hewitt, Chad L.;Campbell, Marnie L.; Schaffelke, Britta (2007) Introductions of seaweeds: accidental transfer pathways and mechanisms., Botanica Marina 50: 326-337

Hiebert, Laurel Sky Vieira, Edson A. Dias, Gustavo M. Tiozzo, Stefano Brown, Federico D. (2019) Colonial ascidians strongly preyed upon, yet dominate the substrate in a subtropical fouling community, Proceedings of the Royal Society of London. Series B 286(20190396.): Published online
http://dx.doi.org/10.1098/rspb.2019.0396

Hill, Steven R. (1996) The flora of Latimer Point and vicinity, New London County, Connecticut, Rhodora 98(894): 180-216

Hillson, Charles J. (1976) Codium invades Virginia waters, Bulletin of the Torrey Botanical Club 103(6): 266-267

Hopkins, C.C.E. (2002) Invasive aquatic species of Europe: Distribution, impacts, and management, Kluwer Academic Publishers, <missing place>. Pp. 240-253

Hubbard, C. B.; Garbary, D. J. (2002) Morphological variation of Codium fragile (Chlorophyta) in eastern Canada, Botanica Marina 45: 476-485

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

Josselyn, Michael N.; West, John A. (1985) The distribution and temporal dynamics of the estuarine macroalgal community of San Francisco Bay, Hydrobiologia 129: 132-152

Karez, Rolf; Ludynia, Katrin (2003) Niche differentiation in habitat and current preference corroborates taxonomic distinction of Jassa falcata and Jassa marmorata (Amphipoda, Crustacea), Estuarine, Coastal and Shelf Science 58: 279-292

Kattler, Kiara R. ; Oishi, Elizabeth M. ; Lim, Em G.; Watkins, Hannah V. ; Côte, Isabelle M. (2023) Functional responses of male and female European green crabs suggest potential sex-specific impacts of invasion, PeerJ 11(e15424): Published online
DOI 10.7717/peerj.15424

Kelly, Jennifer R.; Scheibling, Robert E.; Balch, Toby (2011) Invasion-mediated shifts in the macrobenthic assemblage of a rocky subtidal ecosystem, Marine Ecology Progress Series 437: 69-78

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>

Keppel, E. ; Ruiz, G. M.; Tovar–Hernández, M. A. (2020) Re-description of Parasabella fullo (Grube, 1878) (Polychaeta: Sabellidae) and diagnostic characteristics for detection in California, European Journal of Zoology 87(1): 105-115
DOI: 10.1080/24750263.2020.1721578

Kim, Kwang Young; Garbary, David J. (2007) Photosynthesis in Codium fragile (Chlorophyta) from a Nova Scotia estuary: responses to desiccation and hyposalinity, Marine Biology 151: 99-107

Krumhansl, Kira A.; Scheibling, Robert E. (2011) Detrital production in Nova Scotian kelp beds: patterns and processes, Marine Ecology Progress Series 421: 67-82

Krumhansl, Kira A.; Scheibling, Robert E. (2012) Detrital subsidy from subtidal kelp beds is altered by the invasive green alga Codium fragile ssp. fragile, Marine Ecology Progress Series 456: 73-85

Kusakina, Jelena; Snyder, Marlene; David N.; Dadswell, Kristie; Dadswell, Mike J. (2006) Morphological and molecular evidence for multiple invasions of Codium fragile in Atlantic Canada, Botanica Marina 49: 1-9

Levin, Phillip S.; Coyer, James A.; Petrik, Rachel and Good,Thomas P. (2002) Community-wide effects of nonindigenous species on temperate rocky reefs., Ecology 83(11): 3182-3193

Lewis, J. A. (1999) Center For Research on Introduced Marine Pests,CSIRO Marine Research, Center For Research on Introduced Marine Pests,CSIRO Marine Research, Hobart, Tasmania. Pp. 61-87

Lipej, L.; Mavric, B.; Orlando-Bonaca, M.; Malej, A. (2012) State of the art of the marine non-indigenous flora and fauna in Slovenia, Mediterranean Marine Science 13(2): 243-249

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

Loosanoff, Victor L. (1975) Comment. Introduction of Codium in New England waters, Fishery Bulletin 73(1): 215-218

Loveland, Robert E.; Shafto, Sylvia S. (1984) Fouling Organisms, In: Kennish, Michael J., and Lutz, Richard A.(Eds.) Ecology of Barnegat Bay, New Jersey.. , Berlin. Pp. 226-20

Lyons, Devin A.; Scheibling, Robert E. (2008) Context-dependent survival of the invasive seaweed Codium fragile ssp. tomentosoides in kelp bed and urchin barren habitats off Nova Scotia., Aquatic Biology 2: 17-27

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

Madariaga, David Jofré; Conzález, María Teresa; Dias Bórquez, Christopher; Macaya, Erasmo C.; Harrod, Chris; Thiel. Martin (2023) , None <missing volume>: Published online

Madariaga, David Jofré; González, María Teresa; Bórquez, Christopher Días; Macaya, Erasmo C.'; Harrod, Chris; Thiel, Martin (2023) Successful intertidal colonization of the invasive macroalga Codium fragile near its equatorial/warm range limit in the South‑East Pacific, Biological Invasions 25: 1859–1879
https://doi.org/10.1007/s10530-023-03015-5

Madariaga, David Jofre; Rivadeneira, Marcelo M.; Tala, Fadia; Thiel, Martin (2014) Environmental tolerance of the two invasive species Ciona intestinalis and Codium fragile: their invasion potential along a temperate coast, Biological Invasions 16(12): 2507-2527

Maggi, E. and 11 authors (2015) Ecological impacts of invading seaweeds: a meta-analysis of their effects at different trophic levels, Diversity and Distributions 21: 1-12

Malinowski, K. C.; Ramus, J. (1973) Growth of the green alga Codium fragile in a Connecticut estuary, Journal of Phycology 9: 102-110

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>

Mastrotaro, Francesco; Petrocelli, Antonella; Cecere, Ester; Matarese, Alfonso (2004) Nonidigenous species settle down in the Taranto Seas, Biogeographia 25: 47-58

Matheson, Kyle; McKenzie, Cynthia H.; Sargent, Philip; Hurley, Mike; Wells, Terri (2014) Northward expansion of the invasive green algae Codium fragile spp. fragile (Suringar) Hariot, 1889 into coastal waters of Newfoundland, Canada, BioInvasions Records 3(3): In press

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

Mathieson, Arthur C. ; Dawes, Clinton J.; Hehre, Edward J.; Harris, Larry G. (2010) Floristic Studies of Seaweeds from Cobscook Bay, Maine, Northeastern Naturalist 16(Mongraph 5): 1-48

Mathieson, Arthur C., Hehre, Edward J., Hambrook, Julie, Gerweck, James (1996) A comparison of insular seaweed floras from Penobscot Bay, Maine, and other northwest Atlantic Islands., Rhodora 98(896): 309-418

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.; Hehre, Edward J.; Harris, Larry G. (2009) Floristic studies of seaweeds from Cobscook Bay, Maine, Northeastern Naturalist 16(Monograph 5): 1-48

Mathieson, Arthur C.; Dawes, Clinton J.; Anderson, Michelle L.; Hehre, Edward J. (2001) Seaweeds of the Brave Boat Harbor salt marsh and adjacent open coast of southern Maine., Rhodora 103(913): 1-46

Mathieson, Arthur C.; Dawes, Clinton J.; Harris, Larry; Hehre, Edward J. (2003) Expansion of the Asiatic Green alga Codium fragile subsp. tomentosoides in the Gulf of Maine., Rhodora 105(921): 1-53

Mathieson, Arthur C.; Hehre, Edward J.; Dawes, Clinton J.; Hehre, Edward J. (2008) An historical comparison of seaweed populations from Casco Bay, Maine, Rhodora 110(941): 1-102

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

McDonald, Justin I.; Huisman, John M.; Hart, Felicity N.; Dixon, Rainbo R.M. Lewis, John A. (2015) The first detection of the invasive macroalga Codium fragile subsp. fragile (Suringar) Hariot in Western Australia, BioInvasions Records 4: In press

McNair, N. and Smith, Matt (1999) Investigations into treatments to conrol fouling organisms affecting oyster production., Journal of Shellfish Research 18: 331

Mead, A.; Carlton, J. T.; Griffiths, C. L. Rius, M. (2011b) Introduced and cryptogenic marine and estuarine species of South Africa, Journal of Natural History 39-40: 2463-2524

Meinesz, Alexandre (2007) Methods for identifying and tracking seaweed invasions., Botanica Marina 50: 373-384

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

Miller, Douglas; Brown, Jill. (2005) Maryland’s Coastal Bays: Ecosystem Health Assessment., Maryland Department of Natural Resources., Annapolis. Pp. 8/100-8/105

Miller, Kathy Ann (2004) California's non-native seaweeds, Fremontia 32(1): 10-15

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

Mineur, Frédéric and 12 authors (2015) European seaweeds under pressure: Consequences for communities and ecosystem functioning, Journal of Sea Research 98: 91-108

Mineur, Frederic and 5 authors (2012) Changing Coasts: Marine aliens and artificial structures, Oceanography and Marine Biology, an Annual Review 50: 189-234

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



Morton, B.; Britton, J.C. (2000) Origins of the Azorean intertidal biota: the significance of introduced species, survivors of chance events., Arquipelago. Life and Marine Sciences supplement 2 (part A): 29-51

Nall, Christopher R.; Guerin, Andrew J.; Cook, Elizabeth J. (2015) Rapid assessment of marine non-native species in northern Scotland and a synthesis of existing Scottish records, Aquatic Invasions 10(1): 107–121
http://dx.doi.org/10.3391/ai.2015.10.1.11

Neill, P. E.; Alcalde, O.; Faugeron, S.; Navarrete S. A.; Correa, J. A. (2006) Invasion of Codium fragile ssp. tomentosoides in northern Chile: A new threat for Gracilaria farming, Aquaculture 259: 202-210



NIMPIS (National Introduced Marine Pest Information System). 1998-2015 NIMPIS (National Introduced Marine Pest Information System). <missing URL>



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

Otero, M.; Cebrian, E.; Francour, P.; Galil, B.; Savini, D. (2013) <missing title>, International Union for Conservation of Nature, Malaga, Spain. Pp. 136

Ozturk, Bayram (2002) Invasive Aquatic Species of Europe. Distribution, impacts, and management., Kluwer Academic Publishers, Drodrecht, Netherlands. Pp. 337-340

Pedersen, A.; Kraemer, G.; Yarish, C. (2008) Seaweed of the littoral zone at Cove Island in Long Island Sound: annual variation and impact of environmental factors, Journal of Applied Phycology 20: 869-882

Pederson, Judith and 6 authors (2011) Climate change and non-native species in the North Atlantic, ICES Cooperative Reseach Report 310: 174-190

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

Percy, J. A. 2003 Alien Invasions: introduced species to the Bay of Fundy and environs. <missing URL>



Pérez-Cirera, José Luis; Cremades, Javier; Bárbara, Ignacio (1989) Precisiones sistemáticas y sinecológicas sobre algunas algas nuevas para Galicia o para las costas Atlánticas de la península Ibérica., Anales del Jardin Botanico de Madrid 46(1): 35-45

Petrocelli, Antonella; Cecere, Ester (2015) Biological Invasions in Changing Ecosystems: Vectors, Ecological Impacts, Management and Predictions, de Gruyter, Berlin, Germany. Pp. 253-275

Prince, J. S.; LeBlanc, W. G. (1992) Comparative feeding preference of Strongylocentrotus droebachiensis (Echinoidea) for the invasive seaweed Codium fragile ssp. tomentosoides and four other seaweeds, Marine Biology 113: 159-163

Prince, Jeffrey S. (1988) Sexual reproduction in Codium fragile spp. tomentosoides (Chlorophyceae), Journal of Phycology 24(112-114): <missing location>

Prince, Jeffrey S.; Trowbridge, Cynthia D. (2004) Reproduction in the green macroalga Codium (Chlorophyta): characterization of gametes, Botanica Marina 47: 461–470

Provan, Jim; Booth, David; Todd, Nicola P. ; Beatty, Gemma E.; Maggs, Christine A. (2008) Tracking biological invasions in space and time: elucidating the invasive history of the green alga Codium fragile using old DNA., Diversity and Distributions 39: 43-50

Ramus, Aaron P.; Long, Zachary T. (2016) Producer diversity enhances consumer stability in a benthic marine community, Journal of Ecology 104: 572-579

Ramus, J. (1971) Codium: the invader, Discovery 6(2): 59-68

Ramus, J. (1972) Differentiaon of the green alga Codium fragile, American Journal of Botany 59(5): 478-482

Ribeiro, Romeu S.; Mata, Ana M. T. ; Salgado, Ricardo; Gandra, Vasco; Afonso, Inês; Galhanas, Dina; Dionísio, Maria Ana; Chainho, Paula (2023) Undetected non-indigenous species in the Sado estuary (Portugal), a coastal system under the pressure of multiple vectors of introduction, Journal of Coastal Conservation 27(53): Published online
https://doi.org/10.1007/s11852-023-00979-3

Ribera, Maria A. (1994) <missing title>, European Commission, Brussels. Pp. 37-43

Ribera, Maria Antonia, Boudouresque, Charles-Francois (1995) Introduced marine plants, with special reference to macroalgae: mechanisms and impact., Progress in Phycological Research 11: 187-268

Ruiz, G.M.; Fofonoff, P. W.; Carlton, J. T.; Wonham, M. J.; Hines, A. H. (2000) Invasion of coastal marine communities in North America: Apparent patterns, processes, and biases., Annual Review of Ecology and Systematics 31: 481-531

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>

Saunders, Gary W.; Hawkins, Nick; Wilkin, Sarah (2013) A survey of Sam Orr’s Pond (New Brunswick, Canada) uncovers the invasive green alga Codium fragile (Chlorophyta) and the orange-striped green anemone Diadumene lineata (Cnidaria), first records for the Bay of Fundy and Canada, respectively, Bioinvasions Records 2: in press

Schaffelke, Britta; Deane, David (2005) Desiccation tolerance of the introduced marine green alga Codium fragile ssp. tomentosoides – clues for likely transport vectors?, Biological Invasions 7: 557-565

Schaffelke, Britta; Smith, Jennifer E.; Hewitt, Chad L. (2006) Introduced macroalgae- a growing concern, Journal of Applied Phycology 18: 529-541.

Scheibling, R. E.; Anthony, S. X (2001) Feeding, growth and reproduction of sea urchins (Strongylocentrotus droebachiensis) on single and mixed diets of kelp (Laminaria spp.) and the invasive alga Codium fragile ssp. tomentosoides., Marine Biology 139: 139-146

Scheibling, Robert E.; Lyons, Devin A.; Sumi, Catherine B.T. (2008) Grazing of the invasive alga Codium fragile ssp. tomentosoides by the common periwinkle Littorina littorea: Effects of thallus size, age and condition., Journal of Experimental Marine Biology and Ecology 355: 103-113

Scheibling, Robert E.; Gagnon, Patrick (2006) Competitive interactions between the invasive green alga Codium fragile ssp. tomentosoides and native canopy-forming seaweeds in Nova Scotia (Canada), Marine Ecological Progress Series 325: 1-14

Schmidt, Allison L.; Scheibling, Robert E. (2005) Population dynamics of an invasive green alga, Codium fragile subsp. tomentosoides, in tidepools on a rocky shore in Nova Scotia, Canada, Ecoscience 12(3): 403-412

Schmidt, Allison L.; Scheibling, Robert E. (2006) A comparison of epifauna and epiphytes on native kelps (Laminaria species) and an invasive alga (Codium fragile ssp. tomentosoides) in Nova Scotia, Canada, Botanica Marina 49: 315-330

Schneider, Craig W.; Searles, Richard B. (1991) Seaweeds of the Southeastern United States Cape Hatteras to Cape Canaveral, Duke University Press, Durham. Pp. <missing location>

Searles, R. B.; Hommersand, M. H.; Amsler, C. D. (1984) The occurrence of Codium fragile subsp. tomentosoides and C. taylorii (Chlorophyta) in North Carolina, Botanica Marina 27: 185-187

Silva, Paul C. (1955) The dichotomous species of Codium in Britain, Journal of the Marine Biological Association of the United Kingdom 34: 565-577

Silva, Paul C. (1979) San Francisco Bay Bay: The urbanized estuary; Investigations into the Natural History of San Francisco Bay and Delta With Reference to the Influence of Man, Pacific Division of the American Association for the Advancement of Science, San Francisco. Pp. 287-346

Silva, Paul C.; Menez, Ernani G.; Moe, Richard L. (1987) Catalog of the benthic marine algae of the Philippines, Smithsonian Contributions to the Marine Sciences 27: 1-179

Simard, Par Nathalie; McKindsey, Chris; Archambault, Philippe; Cyr, Charley (2005) Decouverte d'especes marine envahissantes aux iles de la Madeleine., Naturaliste Canadienne 129(2): 62-64

South, G. Robin; Tittley, Ian (1986) <missing title>, Huntsman Marine Laboratory and British Museum (Natural History), St. Andrews, New Brunswick, and London. Pp. <missing location>

Spano, Carlos Spano; Häussermann, Vreni (2017) Anthopleura radians, a new species of sea anemone (Cnidaria: Actiniaria: Actiniidae) from northern Chile, with comments on other species of the genus from the South Pacific Ocean, Biodiversity and Natural History 3: 1-11

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

Streftaris, N.; Zenetos, A. (2006) Alien marine species in the Mediterranean - The 100 ‘worst invasives’ and their impact., Mediterranean Marine Science 7(1): 87-118

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

Taylor, Jonathan E.; Moul, Edwin T.; Loveland, Robert E. (1969) New records and rare benthic marine algae from New Jersey., Bulletin of the Torrey Botanical Club 96(3): 372-378

Thomsen, M. S.; McGlathery, J. (2007) Stress tolerance of the invasive macroalgae Codium fragile and Gracilaria vermiculophylla in a soft-bottom turbid lagoon., Biological Invasions 9: 499-513

Thomsen, M.S.; McGlathery, K. (2006) Effects of accumulations of sediments and drift algae on recruitment of sessile organisms associated with oyster reefs., Journal of Experimental Marine Biology and Ecology 328: 22-34

Thomsen, Madds, S. (2004) <missing title>, Ph.D. Dissertation, University of Virginia, Charlottesville. Pp. <missing location>

Thomsen, Mads S. ; Wernberg, Thomas ; Tuya, Fernando; Silliman, Brian R. (2009) Evidence for impacts of nonindigenous macroalgae: a meta-analysis of experimental field studies, Journal of Phycology 35: 812-819

Thomsen, Mads S.; Wernberg, Thomas; Stæhr, Peter;Krause-Jensen, Dorte; Risgaard-Petersen, Nils; Silliman, Brian R. (2007) Alien macroalgae in Denmark - a broad-scale national perspective., The Korean Journal of Systematic Zoology 3: 61-72

Thorarinsdottir, Gudrun G.; Gunnarsson, Karl; Gíslason, Ó. Sindri (2014) Marine invasive species in the Arctic, Nordic Council of Ministers, Copenhagen, Denmark. Pp. 83-109

Tittley, Ian; Neto, Ana I. (2005) The marine algal (seaweed) flora of the Azores: additions and amendments, Botanica Marina 48: 248-255

Trowbridge, C. D. (1996) Introduced versus native species of Codium fragile: how distinctive is the invasive subspecies tomentosoides?, Marine Biology 126: 193-204

Trowbridge, Cynthia D. (1995) Establishment of the green alga Codium fragile ssp. tomentosoides on New Zealand rocky shores: current distribution and invertebrate grazers, Journal of Ecology 83: 949-965

Trowbridge, Cynthia D. (1997) Dietary induction of opisthobranch morphology: Placida dendritica (Alder & Hancock, 1843) on different green algal diets, Journal of Molluscan Studies 63: 29-38

Trowbridge, Cynthia D. (1998) Ecology of the green macroalga Codium fragile (Suringar) Hariot: invasive and non-invasive subspecies., Oceanography and Marine Biology, an Annual Review 36: 1-64

Trowbridge, Cynthia D. (2004) Emerging associations on marine rocky shores: specialist herbivores on introduced macroalgae, Journal of Animal Ecology 73: 294-308

Trowbridge, Cynthia D.; Farnham, William F. (2009) Regional comparisons of Codium (Chlorophyta) assemblages in the northern versus southern English Channel, Journal of the Marine Biological Association of the United Kingdom 89(2): 255-263

Trowbridge, Cynthia, D.; Todd, Christopher (2001) Host-plant change in marine specialist herbivores: ascoglossan sea slugs on introduced macroalgae, Ecological Monographs 71(2): 219-243

Trowbridge, Cynthis D. (2001) Coexistence of introduced and native congeneric algae: Codium fragile and Codium tomentosum on Irish rocky intertidal shores, Journal of the Marine Biological Association of the United Kingdom 81: 931-937

Tsiamis, Konstantinos; Economou-Amilli, Athena; Katsaros, Christos; Panayotidis, Panayotis (2013) First account of native and alien macroalgal biodiversity at Andros Island (Greece, Eastern Mediterranean), Nova Hedwigia 97(1-2): 209-224

Tsiamis, Konstantinos; Panayotidis, Panayotis (2007) Occurrence of Codium fragile subsp. tomentosoides (van Goor) P.C. Silva (Chlorophyta: Bryopsidophyceae: Bryopsidales: Codiaceae) in Greece., Memoirs of the Museum of Victoria 2(1): 74-76

Valentine, Joseph P.; Magierowski, Regina H.; Johnson, Craig R. (2007) Mechanisms of invasion: establishment, spread and persistence of introduced seaweed populations., Botanica Marina 50: 351-360

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

Verlaque, Marc (1994) Inventaire des plantes introduite en Mediterranee: origines et repercussions sur l'environnment et les activites humaines, Oceanologica Acta 17(1): 1-23

Villalard-Bohnsack, Martine (1995) Illustrated key to the seaweeds of New England, Rhode Island Natural History Survey, Kingston. Pp. <missing location>

Villalard-Bohnsack, Martine, Peckol, Paulette, Harlin, Marilyn M. (1988) Freshwater and Marine Plants of Rhode Island, Kendall/Hunt Publishing Company, Dubuque, Iowa. Pp. 101-118

Villaseñor-Parada, Cristóbal; Macaya, Erasmo C.; Jara-Rojas, Luis M.; Neill, Paula E. (2013) [Spatial and temporal variation in the production of gametangia in the exotic macroalga Codium fragile subsp. tomentosoides (Chlorophyta: Bryopsidales) in the subtidal of Caldera, Chile], Revista de Biología Marina y Oceanografía 48(1): 213-218

Walker, D. I., Kendrick, G. A. (1998) Threats to macroalgal diversity: Marine habitat destruction and fragmentation, pollution and introduced species, Botanica Marina 41: 105-112

Waser, Andreas M.; Lackschewitz, Dagmar; Knol, Jeffrey; Reise, Karsten; Wegner, K. Mathias; Thieltges, David W. (2020) Spread of the invasive shell-boring annelid Polydora websteri (Polychaeta, Spionidae) into naturalised oyster reefs in the European Wadden Sea, Marine Biodiversity 63: <missing location>

Wassman, Robert, Ramus, Joseph (1973) Seaweed invasion, Natural History 82(10): 24-36

Watanabe, Shana; Metaxas, Anna; Scheibling, Robert E. (2009) Dispersal potential of the invasive green alga Codium fragile ssp. fragile, Journal of Experimental Marine Biology and Ecology 381: 114-125

Watanabe, Shana; Scheibling, Robert E.; Metaxas, Anna (2010) Contrasting patterns of spread in interacting invasive species: Membranipora membranacea and Codium fragile off Nova Scotia, Biological Invasions 13: 2329-2342

Wilson, Kristen L.; Kay, Lauren M.; Schmidt, Allison L.; Lotze, Heike K. (2015) Effects of increasing water temperatures on survival and growth of ecologically and economically important seaweeds in Atlantic Canada: implications for climate change, Marine Biology 162: 2431-2444

Wiltshire, K.; Rowling, K.; Deveney, M. (2010) <missing title>, South Australian Research and Development Institute, Adelaide. Pp. 1-232

Zaitsev, Yuvenali; Ozturk, Bayram (2001) <missing title>, Turkish Marine Research Foundation Publication, <missing place>. Pp. 1-265

---