Description
Other Taxonomic Categories -A number of subspecies of Codium fragile are known, and are found in most temperate regions of the world. Codium fragile spp. fragile (Dead Man's Fingers) is believed to have originated in the Northwest Pacific in waters around Japan, where the species and subspecies were first described (Guiry et al. 2009). Codium f. ssp. fragile, formerly known as C. f. ssp. tomentosoides (Provan et al. 2007; Guiry et al. 2009) is the most widely distributed "weed" species, but C. f. ssp. atlanticum and C. f. ssp. scandinavicum are also probably introduced to Europe (Silva 1955, 1957; cited by Trowbridge 1996). Subspecies of C. fragile are native to areas (Pacific Coast, New Zealand) which have been invaded by C. f. ssp. fragile.
Potentially Misidentified Species - No native species of Codium are known from our area or elsewhere in the Northwest Atlantic (South and Tittley 1986); 4 species are found in NC and southwards (C. carolinianum; C. decorticatum; C. isthmocladium; C. taylori) (Schneider and Searles 1992).
Taxonomy
| Kingdom | Phylum | Class | Order | Family | Genus |
|---|---|---|---|---|---|
| Protista | Chlorophyta | Ulvophyceae | Caulerpales | Codiaceae | Codium |
Synonyms
Invasion History
Chesapeake Bay Status
| First Record | Population | Range | Introduction | Residency | Source Region | Native Region | Vectors |
|---|---|---|---|---|---|---|---|
| 1976 | Established | Expanding | Introduced | Regular Resident | Western Atlantic | Western Pacific | Shipping(Fouling Community, Ballast Water),Fisheries(Packing Material-Bait); Oysters-Accidental); Natural Dispersal(Natural Dispersal) |
History of Spread
Codium fragile spp. fragile (Dead Man's Fingers) is believed to have originated in the Northwest Pacific in waters around Japan. In Europe, it was first collected in Holland in 1900 and in England in 1939 and has spread extensively along Northeast Atlantic coastlines (Farnham 1980) from Iceland and Norway to France (South and Tittley 1986). It was first collected in the Mediterranean in France in1950 (Verlaque 1994), and has spread east to the straits of Messina, Italy, and Libya (Ribera and Boudouresque 1995) It has also been introduced to the Azores, San Francisco Bay, Australia, and New Zealand (Cohen and Carlton 1995; Hewitt et al. 2000; Trowbridge 1995; (Cardigos et al. 2006)
Codium fragile spp. fragile 's first appearance in Northwest Atlantic waters was in Long Island Sound in 1957. Its introduction has been attributed to ship fouling, or to 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. spp. fragile, The spotty nature of its spread in New England waters suggests a diversity of transport mechanisms.
Northwest Atlantic records are summarized below:
Nova Scotia - Codium fragile spp. 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. It may have been brought with transplanted shellfish, as packing material for bait or shellfish, or by Gulf Stream eddies from New England (Bird et al. 1993). In 1996, it was collected for the first time in the Gulf of St. Lawrence, in Caribou Harbour, on the Northumberland Straits (Garbary et al. 1997).
Gulf of Maine - Codium fragile spp. fragile was first collected north of Cape Cod in Boothbay Harbor ME 1964, but its spread in Cape Cod Bay occurred later. It reached the Cape Cod Canal by 1969, and successively was found in: Barnstable Harbor MA (1972) Wellfleet Harbor MA (1974) on the western side of the Bay, Duxbury Bay (1981), and Provincetown (1981). By 1983, it was collected at Appledore Island, Isles of Shoals NH-M ,(Carlton and Scanlon 1985). However, it was present to the north, at Bar Harbor ME by the late 1970's or early 1980's (Bird et al. 1993).
Long Island Sound, Southern New England Bays, and Sounds - Codium fragile spp. fragile was first collected in East Marion NY on the Eastern Fork, in Gardiners's and Peconic Bays in 1957 (Bouck and Morgan 1957). It reached the CT side by 1961; and RI by 1962-68, and then was 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 rhe south side of Cape Cod (Carlton and Scanlon 1985).
NJ - By 1966, Codium fragile spp. fragile was found in Barnegat Bay (Carlton and Scanlon 1985).
Chesapeake Bay Region - Codium fragile spp. fragile was first collected in the region in Assateague Channel VA in 1976. This colony was probably 1-2 years old (Hillson 1976). There are few recent records, but it was seen washed up on beach on Poquoson Neck VA, on the western side of the lower Bay in 1995. Codium fragile spp. fragile was not found on beaches of the Eastern Shore (Godwin 1997, personal communciation). It was found at Cape Henry VA in 1999 (Ruiz et al., unpublished data), and is common to abundant in Atlantic coastal bays. It was the fourth most abundant macroalga in Hog Island Bay VA, just north the Chesapeake Bay's entrance (Thomsen 2004).
NC - Codium fragile spp. fragile approached its southernmost limit at Topsail Inlet, south of Cape Hatteras in 1979; Masonboro Inlet 1982, and Beaufort Inlet (1981) (Searles et al. 1984). Its range apparently has not expanded southward since then (Schneider and Searles 1992; Searles 1997).
History References - Bird et al. 1993; Bouck and Morgan 1957; Carlton and Scanlon 1984; Farnham 1980; Garbary et al. 1997; Godwin 1997; Hillson 1976; Schneider and Searles 1991; Searles et al. 1984; Searles 1997; South and Tittley 1986; Thomsen 2003
Invasion Comments
Ecology
Environmental Tolerances
| For Survival | For Reproduction | |||
|---|---|---|---|---|
| Minimum | Maximum | Minimum | Maximum | |
| Temperature (ºC) | -2.0 | 30.0 | 12.0 | 30.0 |
| Salinity (‰) | 18.0 | 42.0 | 36.0 | 22.0 |
| Oxygen | ||||
| pH | ||||
| Salinity Range | poly-eu |
Age and Growth
| Male | Female | |
|---|---|---|
| Minimum Adult Size (mm) | 150.0 | |
| Typical Adult Size (mm) | 200.0 | |
| Maximum Adult Size (mm) | 250.0 | |
| Maximum Longevity (yrs) | 2.5 | |
| Typical Longevity (yrs |
Reproduction
| Start | Peak | End | |
|---|---|---|---|
| Reproductive Season | |||
| Typical Number of Young Per Reproductive Event |
|||
| Sexuality Mode(s) | |||
| Mode(s) of Asexual Reproduction |
|||
| Fertilization Type(s) | |||
| More than One Reproduction Event per Year |
|||
| Reproductive Startegy | |||
| Egg/Seed Form |
Impacts
Economic Impacts in Chesapeake Bay
Problems due to Codium fragile ssp. fragile have not been reported in the Chesapeake Bay region. The alga's salinity requirements would restrict it to the mouth of the Bay proper, but it could reach troublesome densities in the embayments along the Atlantic shore to the north.
Economic Impacts Outside of Chesapeake Bay
Codium fragile ssp. fragile is widely regarded as a troublesome weed in areas where it has been introduced.
Fisheries - As noted under 'Ecological Impacts', C. f ssp. fragile attaches 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 water (Fofonoff personal observation).
Aesthetic - Accumulations of Codium fragile on beaches are unattractive and unpleasant to walk on, perhaps more so than most seaweeds (Fofonoff personal observation).
References - Ramus 1971
Ecological Impacts on Chesapeake Native Species
Impacts of Codium fragile ssp. fragile (Dead Man's Fingers) are probably small in the Chesapeake Bay region, since this seaweed appears to be rare, with a restricted distribution (Hillson 1976; Godwin 1997).
Competition - Codium fragile ssp. fragile' advantages over native seaweeds include rapid growth and longevity, since most native plants are annuals. It rapidly responds to added nutrients, giving it an advantage in eutrophic areas. It appears to do best as a colonizer of bare surfaces, and rarely invades areas of established vegetation (Ramus 1971). In Hog Island Bay, Virginia, C. f. ssp. 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).
Food/Prey - In regions where invertebrate grazing on it has been studied (New England; New Zealand), C. f. ssp. fragile is eaten by native gastropods, chitons, and echinoids, but is usually not preferred over native seaweeds, including, in New Zealand, native subspecies of C. fragile (Prince and LeBlanc 1992; Trowbridge 1995). A range extension of the ascoglossan sea slug Placida dendritica in New England may have been facilitated by this seaweed's invasion (Clark and Franz 1968), but this opisthobranch feeds on other green algae in addition to C. f. ssp. fragile (Bleakney 1989). In New Zealand it prefers native forms of Codium over C. f. ssp. fragile (Trowbridge 1996).
Predation - Codium fragile ssp. fragile obviously does not eat shellfish, but it can kill them indirectly by attaching to their shells. When this seaweed attaches to Bay Scallops (Argopecten irradians), it interferes with the scallops' swimming movements and its ability to escape predators. Because the tissues of C. f. ssp. fragile become buoyant during periods of photosynthesis, scallops, mussels (Mytilus edulis), and oysters Crassostrea virginica can be dragged by the current or float to the surface and ultimately washed ashore. This appeared to be an important source of shellfish mortality in shallow areas of Long Island Sound. Hence this seaweed is sometimes called 'oyster thief' (Ramus 1971).
Habitat Change - Codium fragile ssp. fragile probably provides cover for some fishes and invertebrates. In southern New England coastal waters, juvenile Black Sea Bass Centropristes striatus are frequently found under stones or shells attached to Codium (Fofonoff 1970-1980s' personal observation). Codium fragile ssp. fragile also affects the composition of beaches by increasing the deposition of stones and shells (with weed attached) (Ben-Avraham 1971).
References - Ben-Avraham 1971; Bleakney 1989; Clark and Franz 1968; Godwin 1997; Hillson 1976; Prince and LeBlanc 1992; Ramus 1971; Trowbridge 1996
Ecological Impacts on Other Chesapeake Non-Native Species
Impacts of Codium fragile ssp. fragile (Dead Man's Fingers) on other introduced seaweed species are probably small, since most of the introduced forms appear to be uncommon. In the Atlantic embayments to the north of the Chesapeake, C. f. ssp. fragile may provide cover to such invertebrates as Carcinus maenas (Green Crab) and Littorina littorea (Common Periwnkle).
References
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: 11-17Bouck, G. B.; Morgan, E. (1957) The occurrence of Codium in Long Island waters, Bulletin of the Torrey Botanical Club 84: 384-387
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
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
Clark, Kerry B.; Franz, David R. (1968) Occurrence of the saccoglossan opisthobranch Hermaea dendritica Alder & Hancock in New England, Veliger 12: 174-175
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, , Washington DC, Silver Spring MD.. Pp.
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
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
1997 conversation with Paul Fofonoff. conversation
2004-2023 AlgaeBase. National University of Ireland Galway--http://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
Hewitt, Chad L.; Campbell, Marnie L. (2000) Historical and Modern Invasions to Port Phillip Bay, Australia: The Most Invaded Southern Embayment?, In: (Eds.) Marine Bioinvasions. , . Pp. 120-126
Hillson, Charles J. (1976) Codium invades Virginia waters, Bulletin of the Torrey Botanical Club 103: 266-267
Loosanoff, Victor L. (1975) Comment. Introduction of Codium in New England waters, Fishery Bulletin 73: 215-218
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
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, J. (1971) Codium: the invader, Discovery 6: 59-68
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
Schneider, Craig W.; Searles, Richard B. (1991) Seaweeds of the Southeastern United States, , Durham. Pp.
Searles, R. B. (1984) Seaweed biogeography of the mid-Atlantic coast of the United States, Helgoländer Meeresuntersuchungen 38: 259-271
March 21, 1997 Introduced seaweeds in Chesapeake Bay, email. Department of Botany, Duke University
South, G. Robin; Tittley, Ian (1986) A checklist and distributional index of the benthic marine algae of the North Atlantic Ocean., , St. Andrews, New Brunswick, and London. Pp.
Thomsen, Madds, S. (2004) Macroalgal distribution patterns and ecological performances in a tidal coastal lagoon, with emphasis on the non-indigenous Codium fragile ssp. tomentosoides., None , Charlottesville. Pp. None
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
Verlaque, Marc (1994) Inventaire des plantes introduite en Mediterranee: origines et repercussions sur l'environnment et les activites humaines, Oceanologica Acta 17: 1-23