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Datasheet

Codium fragile subsp. tomentosoides

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Datasheet

Codium fragile subsp. tomentosoides

Summary

  • Last modified
  • 13 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Preferred Scientific Name
  • Codium fragile subsp. tomentosoides
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Chlorophyta
  •       Class: Bryopsidophyceae
  •         Order: Bryopsidales
  • Summary of Invasiveness
  • C. fragile ssp. tomentosoides is considered as an invasive marine alga as it has the capacity to spread rapidly via asexual reproduction and fragmentation, and has invaded many coastal waters including tho...

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Pictures

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PictureTitleCaptionCopyright
Codium fragile ssp. tomentosoides in natural habitat.
TitleNatural habit
CaptionCodium fragile ssp. tomentosoides in natural habitat.
CopyrightSuchana A. Chavanich
Codium fragile ssp. tomentosoides in natural habitat.
Natural habitCodium fragile ssp. tomentosoides in natural habitat.Suchana A. Chavanich

Identity

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Preferred Scientific Name

  • Codium fragile subsp. tomentosoides

International Common Names

  • English: dead man’s fingers; green fleece

Local Common Names

  • Netherlands: viltwier
  • UK/England and Wales: green sea fingers
  • USA: sputnik weed

Summary of Invasiveness

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C. fragile ssp. tomentosoides is considered as an invasive marine alga as it has the capacity to spread rapidly via asexual reproduction and fragmentation, and has invaded many coastal waters including those of Europe and America. Its spread has also had a negative impact on benthic communities. C. fragile ssp. tomentosoides can become a dominant member in invaded habitats and can alter community composition and function. In addition, it can tolerate wide ranges of temperature and salinity which contribute to it becoming a dominant species when conditions permit.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Chlorophyta
  •             Class: Bryopsidophyceae
  •                 Order: Bryopsidales
  •                     Family: Codiaceae
  •                         Genus: Codium
  •                             Species: Codium fragile subsp. tomentosoides

Notes on Taxonomy and Nomenclature

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The centre of Codium fragile diversity is assumed to be in East Asia (Japan and Korea), where the species is highly variable (Silva, 1955; Provan et al., 2005). In general, C. fragile is considered to have six subspecies, three of which (including C. fragile ssp. tomentosoides) are considered as weeds in several parts of the world (Silva, 1955; 1957; Trowbridge, 1998; 1999). However, in Japan and Korea, C. fragile is considered a single species with no subspecies (Segawa, 1996; Chavanich et al., 2006).

Description

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C. fragile ssp. tomentosoides is a dark green alga, up to 1 m in length. It is composed of dichotomous cylindrical branches, 0.5-1.0 cm in diameter. It is made up of interwoven filaments ending in a layer of swollen utricles that cover the surface of the thallus (Villalard-Bohnsack, 1995). Reproduction of C. fragile ssp. tomentosoides is either by motile unicells or vegetative fragments(Chapman, 1999; Mathieson et al.,2003). C. fragile ssp. tomentosoides also has parthenogenetic gametes that germinate without fertilization (Trowbridge, 1999). During the winter, extensive fragmentations occur as a result of low temperatures which cause thallus constriction and segmentation (Fralick and Mathieson, 1972).

Distribution

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C. fragile ssp. tomentosoides has spread throughout the northern and southern hemispheres including the northeastern and northwestern Atlantic, the Mediterranean, Australia, New Zealand, the eastern central Pacific, and southeastern Pacific. In east Asia (Japan and Korea), where Codium fragile is a native species, its spread is restricted to areas where water temperatures are between 10-20°C (Lee and Kang, 1986; Segawa, 1996). However, when invading new habitats, C. fragile ssp. tomentosoides can withstand temperatures as low as -2°C (Fralick and Mathieson, 1972).

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Sea Areas

Atlantic, NortheastWidespreadIntroduced Invasive Trowbridge, 1999; Provan et al., 2005; Guiry and Guiry, 2007
Atlantic, NorthwestWidespreadIntroduced Invasive Trowbridge, 1999; Hubbard and Garbary, 2002; Begin and Scheibling, 2003; Provan et al., 2005
Atlantic, SoutheastWidespreadIntroduced Invasive Begin and Scheibling, 2003
Atlantic, Western CentralWidespreadIntroduced Invasive Provan et al., 2005
Indian Ocean, EasternWidespreadIntroduced Invasive Trowbridge, 1999
Mediterranean and Black SeaWidespreadIntroduced Invasive Trowbridge, 1999; Provan et al., 2005; Guiry and Guiry, 2007
Pacific, Eastern CentralWidespreadIntroduced Invasive Trowbridge, 1999
Pacific, NorthwestLocalisedNative Not invasive Provan et al., 2005; Chavanich et al., 2006
Pacific, SoutheastWidespreadIntroduced Invasive Provan et al., 2005
Pacific, SouthwestWidespreadIntroduced Invasive Trowbridge, 1998; Begin and Scheibling, 2003

Asia

JapanPresentPresent based on regional distribution.
-HokkaidoLocalisedNative Not invasive Provan et al., 2005
-HonshuLocalisedNative Not invasive Provan et al., 2005; Chavanich et al., 2006
Korea, Republic ofLocalisedNative Not invasive Chavanich et al., 2006
TurkeyWidespreadIntroduced Invasive Guiry and Guiry, 2007

Africa

AlgeriaWidespreadIntroduced Invasive Guiry and Guiry, 2007
South AfricaWidespreadIntroduced Invasive Begin and Scheibling, 2003
TunisiaWidespreadIntroduced Invasive Guiry and Guiry, 2007

North America

CanadaPresentPresent based on regional distribution.
-New BrunswickWidespreadIntroduced Invasive Hubbard and Garbary, 2002; Begin and Scheibling, 2003
-Nova ScotiaWidespreadIntroduced Invasive Chapman, 1999; Hubbard and Garbary, 2002; Begin and Scheibling, 2003
-Prince Edward IslandWidespreadIntroduced Invasive Hubbard and Garbary, 2002; Begin and Scheibling, 2003
USAPresentPresent based on regional distribution.
-CaliforniaWidespreadIntroduced Invasive Trowbridge, 1999First found in 1977
-ConnecticutWidespreadIntroduced Invasive Carlton and Scanlon, 1985; Begin and Scheibling, 2003First found in Fishers Island Sound in 1961
-DelawareWidespreadIntroduced Invasive Trowbridge, 1999
-MaineWidespreadIntroduced Invasive Carlton and Scanlon, 1985; Begin and Scheibling, 2003First found in Boothbay Habour in 1964
-MassachusettsWidespreadIntroduced Invasive Carlton and Scanlon, 1985; Begin and Scheibling, 2003First found on south shore of Cape Cod in 1961
-New HampshireWidespreadIntroduced Invasive Carlton and Scanlon, 1985; Provan et al., 2005First found in Isles of Shoals, Appledore Island in 1983
-New JerseyWidespreadIntroduced Invasive Carlton and Scanlon, 1985; Trowbridge, 1999First found in Barnegat Bay in 1966
-New YorkWidespreadIntroduced Invasive Carlton and Scanlon, 1985; Begin and Scheibling, 2003First found in Long Island Sound in 1957
-North CarolinaWidespreadIntroduced Invasive Carlton and Scanlon, 1985; Provan et al., 2005First found in Onslow Bay between Cape Fear and Cape Lookout in 1979
-Rhode IslandWidespreadIntroduced Invasive Carlton and Scanlon, 1985; Trowbridge, 1999First found in Narragansett Bay between 1962 and 1968
-VirginiaWidespreadIntroduced Invasive Carlton and Scanlon, 1985; Trowbridge, 1999First found in Assateague Channel in 1976

South America

ChileWidespreadIntroduced Invasive Neill et al., 2003; Provan et al., 2005First record in Northern Chile in 1998

Europe

DenmarkWidespreadIntroduced Invasive Trowbridge, 1999First recorded in 1919
FranceWidespreadIntroduced Invasive Provan et al., 2005; Guiry and Guiry, 2007
GreeceWidespreadIntroduced Invasive Provan et al., 2005; Tsiamis and Panayotidis, 2007First found in Maliakos Gulf in 1992
IrelandWidespreadIntroduced Invasive Trowbridge, 1999; Provan et al., 2005; Guiry and Guiry, 2007First recorded in 1941
ItalyWidespreadIntroduced Invasive Guiry and Guiry, 2007
NetherlandsWidespreadIntroduced Invasive Trowbridge, 1999; Provan et al., 2005First recorded in ca. 1900
NorwayWidespreadIntroduced Invasive Trowbridge, 1999First recorded in 1946
PortugalWidespreadIntroduced Invasive Guiry and Guiry, 2007
SloveniaWidespreadIntroduced Invasive Provan et al., 2005
SpainWidespreadIntroduced Invasive Provan et al., 2005; Guiry and Guiry, 2007
SwedenWidespreadIntroduced Invasive Trowbridge, 1999First recorded in 1933
UKWidespreadIntroduced Invasive Trowbridge, 1999; Provan et al., 2005; Guiry and Guiry, 2007First recorded in England in 1939, and first recorded in Scotland in 1953

Oceania

AustraliaPresentPresent based on regional distribution.
-New South WalesWidespreadIntroduced Invasive Trowbridge, 1999
-South AustraliaWidespreadIntroduced Invasive Trowbridge, 1999
-TasmaniaWidespreadIntroduced Invasive Trowbridge, 1999
-VictoriaWidespreadIntroduced Invasive Trowbridge, 1999First found in 1995 at Corner Inlet
-Western AustraliaWidespreadIntroduced Invasive Trowbridge, 1999
New ZealandWidespreadIntroducedTrowbridge, 1999; Begin and Scheibling, 2003first found in Auckland Harbour in 1973

History of Introduction and Spread

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C. fragile ssp. tomentosoides is native to east Asia; the first exotic occurrence being in Holland around 1900, presumably introduced with shellfish (Silva, 1955, 1957). It then spread rapidly across the Mediterranean and Europe during the World War II (Silva, 1955, 1957; Trowbridge and Todd, 1999). It was first observed in Long Island Sound on the northwest Atlantic Coast in 1957 (Bouck and Morgan, 1957). Later in 1977, it was found in San Francisco Bay, California, USA (Silva, 1979). In the southern hemisphere, it was first found in Auckland Harbour, New Zealand in 1973 (Dromgoole, 1975).

Risk of Introduction

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There are three potential vectors for the transoceanic and or transcontinental dispersal of C. fragile ssp. tomentosoides:


1) fouling of the flat oyster Ostrea edulis from Europe

2) fouling of the Japanese or Pacific oyster Crassostrea gigas from Washington and British Columbia

3) fouling of ship’s hulls from Europe (Carlton and Scanlon, 1985).

 

It also can be spread via boating, packing material for fishery products (such as lobsters and bait worms), and water currents (Carlton and Scanlon, 1985).

Habitat

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C. fragile ssp. tomentosoides can be found year around on open coasts, estuaries, tide pools, and intertidal and subtidal zones. It attaches to rocks, shells, or other hard substrates, and is often covered with epiphytic species (Villalard-Bohnsack, 1995). It is also found in sheltered habitats such as bays and harbours (Mathieson et al., 2003). In its native range, C. fragile is a member of the underwater understory assemblage below the dominant canopy species whereas in disturbed or invaded habitats, C. fragile is a dominant canopy species (Chavanich et al., 2006).  In addition, in its native range, it can be found to depths of 18 m, and colonizes novel substrates such as floating docks and piles (Chavanich et al., 2006).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Littoral
Coastal areas Principal habitat Harmful (pest or invasive)
Coastal areas Principal habitat Natural
Coastal areas Principal habitat Productive/non-natural
Mud flats Present, no further details Harmful (pest or invasive)
Mud flats Present, no further details Natural
Mud flats Present, no further details Productive/non-natural
Intertidal zone Present, no further details Harmful (pest or invasive)
Intertidal zone Present, no further details Natural
Intertidal zone Present, no further details Productive/non-natural
Salt marshes Present, no further details Harmful (pest or invasive)
Salt marshes Present, no further details Natural
Salt marshes Present, no further details Productive/non-natural
Brackish
Estuaries Present, no further details Harmful (pest or invasive)
Estuaries Present, no further details Natural
Estuaries Present, no further details Productive/non-natural
Marine
 
Benthic zone Principal habitat Harmful (pest or invasive)
Benthic zone Principal habitat Natural
Benthic zone Principal habitat Productive/non-natural

Biology and Ecology

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Provan et al.(2005) reported that in its native range of Japan, there were low levels of genetic variation in C. fragile ssp. tomentosoides, with only four haplotypes present. Two of these haplotypes were also found in populations introduced to other countries from Japan. The authors conclude that there were at least two separate introductions of C. fragile ssp. tomentosoides from Japan.

Reproductive Biology

Sexual, parthenogenetic and vegetative reproduction has been reported in C. fragile ssp. tomentosoides (Fralick and Mathieson, 1972; Chapman, 1999; Trowbridge, 1999). Gametangia containing two types of biflagellate cells have been found, although the function of male gametes is still unclear (Prince, 1988; Trowbridge, 1999).  Bulleri et al. (2007) found that habitat had an influence on the growth and reproduction of C. fragile ssp. tomentosoides.


Vegetative reproduction of C. fragile ssp. tomentosoides occurs through fragmentation of the thallus. Maximum fragmentation of C. fragile ssp. tomentosoides occurs during the winter as the reduced temperature causes constrictions or swellings of the thalli (Fralick and Mathieson, 1972).

 
Physiology and Phenology

Fralick and Mathieson (1973) showed that optimal conditions for net photosynthesis of C. fragile ssp. tomentosoides were between 21-24°C and 900-1100 foot-candles.

Associations


Epiphytes and herbivores can be found associated with Codium fragile both in native and non-native ranges (Mathieson et al., 2003; Chavanich et al., 2006).  The epiphytic red algae Ceramium nodulosum is commonly found on C. fragile ssp. tomentosoides in the New England area, USA (Villalard-Bohnsack, 1995). Trowbridge (1999) found at least 17 species of algae epiphytic on C. fragile ssp. tomentosoides. Several groups of herbivores such as molluscs, crustaceans, echinoderms, and bryozoans use C. fragile ssp. tomentosoides as their habitat and food source (Chavanich and Harris, 2000; Harris and Mathieson, 2000; Harris and Tyrrell, 2001; Scheibling and Anthony, 2001; Trowbridge and Todd, 2001; Chavanich and Harris, 2002; Chavanich and Harris, 2004). However, several studies have showed that C. fragile ssp. tomentosoides is not a preferred food (Prince and Le Blanc, 1992; Trowbridge, 1995; Chavanich and Harris, 2004).

 
Environmental Requirements
 

C. fragile ssp. tomentosoides occurs on exposed coasts with protected bays. It also can be found in areas with extremely oligotrophic to euthrophic nutrients (Chapman, 1999).

 

Climate

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ClimateStatusDescriptionRemark
Cf - Warm temperate climate, wet all year Tolerated Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year
Cs - Warm temperate climate with dry summer Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers
Cw - Warm temperate climate with dry winter Tolerated Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)
Df - Continental climate, wet all year Tolerated Continental climate, wet all year (Warm average temp. > 10°C, coldest month < 0°C, wet all year)
Ds - Continental climate with dry summer Tolerated Continental climate with dry summer (Warm average temp. > 10°C, coldest month < 0°C, dry summers)
Dw - Continental climate with dry winter Tolerated Continental climate with dry winter (Warm average temp. > 10°C, coldest month < 0°C, dry winters)

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
60 50 0 0

Water Tolerances

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ParameterMinimum ValueMaximum ValueTypical ValueStatusLife StageNotes
Depth (m b.s.l.) 1 9 Optimum
Salinity (part per thousand) 30 35 Optimum 12-40 tolerated
Water temperature (ºC temperature) 10 25 Optimum -2-34 tolerated

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Elysia maoria Herbivore Whole plant to genus Trowbridge, 1995
Elysia viridis Herbivore Whole plant to genus Trowbridge and Todd, 2001
Lacuna vincta Herbivore Whole plant not specific Chavanich and Harris, 2004
Placida dendritica Herbivore Whole plant to genus Harris and Mathieson, 2000; Trowbridge, 1995
Strongylocentrotus droebachiensis Herbivore Whole plant not specific Prince and LeBlanc, 1992
Turbo smaragdus Herbivore Whole plant not specific Trowbridge, 1995

Notes on Natural Enemies

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Natural enemies of C. fragile ssp. tomentosoides are usually herbivores such as molluscs, crustaceans, and echinoderms (Harris and Mathieson, 2000; Harris and Tyrrell, 2001; Scheibling and Anthony, 2001; Trowbridge and Todd, 2001; Chavanich and Harris, 2002; Chavanich and Harris, 2004). However, because these herbivores are small and C. fragile ssp. tomentosoides is not a palatable alga (Prince and LeBlanc, 1992; Trowbridge, 1995; Chavanich and Harris, 2004), the impact of herbivores on C. fragile ssp. tomentosoides populations is quite low. One group of snails of the genus Sacoglossa, typically found associated with siphonaceous alga, contain several species that are known to feed on C. fragile (Trowbridge, 1992; 1993; Harris and Mathieson, 2000; Trowbridge and Todd, 2001; Trowbridge, 2002; van Bragt, 2004).

Means of Movement and Dispersal

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C. fragile ssp. tomentosoides can be spread or introduced through natural dispersal such as water currents and by accidental introduction via ship’s hulls and shellfish. It also can be spread via boating, and packing material for fishery products.

There are four major dispersal mechanisms of C. fragile ssp. tomentosoides (from Carlton and Scanlon, 1985):



  1. Dispersal as a ship fouling organism
  2. Transport via attachment to the shells of commercial oysters
  3. Accidental introduction by commercial fishermen who collect Codium with drag nets.
  4. Dispersal with other algae used as packing material for fishery products such as lobsters and bait worms.

  5. Harris and Tyrrell (2001) also report on the possible linkage between the spread of C. fragile ssp. tomentosoides and climate change in the Gulf of Maine, USA.

    Impact Summary

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    CategoryImpact
    Economic/livelihood Negative
    Environment (generally) Negative

    Economic Impact

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    C. fragile ssp. tomentosoides can have an economic impact on shellfish and fishing industries. It can smother mussels and scallops, reduce the biomass of oysters, lift the shellfish off the sea floor, foul the nets, and clog scallop dredges (Trowbridge, 1999). This leads to an increase in labour costs during harvesting and processing associated with the need to remove the algae (Carlton and Scanlon, 1985; Trowbridge, 1999).

    Environmental Impact

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    Impact on Habitats

    Because C. fragile ssp. tomentosoides is an opportunistic alga, its spread can alter the ecosystem by replacing the dominant species such as kelp (Harris and Tyrrell, 2001). In native habitats, C. fragile ssp. tomentosoides not only increases its density in benthic communities but also colonizes novel substrates like floating docks and pilings (Chavanich et al., 2006).

    Impact on Biodiversity
     

    Consequently, changes in community structure and composition, such as that observed in the Gulf of Maine, by C. fragile ssp. tomentosoides have affected other native species by shifting habitat selection and feeding behaviours (Chavanich and Harris, 2004; Harris and Jones, 2005). This introduced alga can also cause the decline of populations of native species (Chavanich and Harris, 2004).

     

    Threatened Species

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    Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
    Lacuna vinctaNo details No detailsUSAChavanich and Harris, 2004
    Laminaria saccharinaNo details No detailsCanada; USAHarris and Tyrrell, 2001; Scheibling and Anthony, 2001

    Social Impact

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    In native ranges in Korea, C. fragile is consumed by local people as soup.

    Risk and Impact Factors

    Top of page Invasiveness
    • Invasive in its native range
    • Proved invasive outside its native range
    • Highly adaptable to different environments
    • Is a habitat generalist
    • Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
    • Pioneering in disturbed areas
    • Long lived
    • Fast growing
    • Has high reproductive potential
    • Reproduces asexually
    • Has high genetic variability
    Impact outcomes
    • Altered trophic level
    • Conflict
    • Damaged ecosystem services
    • Ecosystem change/ habitat alteration
    • Increases vulnerability to invasions
    • Infrastructure damage
    • Modification of natural benthic communities
    • Modification of successional patterns
    • Monoculture formation
    • Negatively impacts aquaculture/fisheries
    • Reduced native biodiversity
    • Threat to/ loss of native species
    Impact mechanisms
    • Competition - monopolizing resources
    • Competition - shading
    • Competition - smothering
    • Fouling
    • Interaction with other invasive species
    • Rapid growth
    Likelihood of entry/control
    • Highly likely to be transported internationally accidentally
    • Difficult to identify/detect as a commodity contaminant
    • Difficult/costly to control

    Similarities to Other Species/Conditions

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    Gross morphological differences among the subspecies of C. fragile are not obvious. However, they can be differentiated on the variation in size and shape of utricles (Silva, 1955, 1957). C. fragile ssp. tomentosoides has utricles with pointed apices and a distinct constriction in the middle (Villalard-Bohnsack, 1995; Trowbridge, 1999).

    Prevention and Control

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    Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

    To date there are no obvious action plans involving the management and eradication of C. fragile ssp. tomentosoides. There are also no plans to restore habitats invaded by this alga. However, Trowbridge (1999) suggests several potential methods of reducing and preventing further spread of C. fragile ssp. tomentosoides:



    1. Interventionist control methods (species reduction or eradication)


      1. Chemical treatment such as using Diquat, Stomp, copper sulphate or sodium hypochlorite

      2. Mechanical control such dredging and dragging

      3. Manual removal

      4. Biological control by using specialized sea slugs (ascoglossans) that feed on Codium species.

    2. Quarantine control methods


      1. Prevent contaminated shellfish from being introduced to new areas

      2. Exclusion of drift thalli to prevent the dispersal of gametes.

      Public awareness campaigns regarding this alga have been carried out in several parts of the USA in the form of brochures and websites.

      References

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      Begin C; Scheibling RE, 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, No. 46:404-412.

      Bouck GB; Morgan E, 1957. The occurrence of Codium in Long Island waters. Bulletin of the Torrey Botanical Club, No. 84:384-387.

      Bragt PHVan, 2004. The sea slugs, Sacoglossa and Nudibranchia (Gastropoda, Opisthobranchia), of the Netherlands. Vita Malacologica, No. 2:3-32.

      Bulleri F; Branca MG; Abbiati M; Airoldi L, 2007. Development of reproductive structures in the introduced green alga, Codium fragile ssp. tomentosoides, in the northern Adriatic Sea. European Journal of Phycology, No. 42:137-144.

      Carlton JT; Scanlon JA, 1985. Progression and dispersal of an introduced alga: Codium fragile ssp. tomentosoides (Chlorophyta) on the Atlantic coast of North America. Botanica Marina, No. 28:155-165.

      Chapman AS, 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, No. 52:277-289.

      Chapman VJ; Chapman DJ, 1980. Seaweeds and their Uses. London, UK: Chapman and Hall.

      Chavanich S; Harris LG, 2000. Marine Bioinvasions [ed. by Pederson J]. USA: Massachusetts Institute of Technology, MA, 157-163.

      Chavanich S; Harris LG, 2002. The influence of macroalgae on seasonal abundance and feeding preference of a subtidal snail, Lacuna vincta (Montagu) (Littorinidae) in the Gulf of Maine. Journal of Molluscan Studies, No. 68:73-78.

      Chavanich S; Harris LG, 2004. Impact of the non-native macroalgae, Codium fragile (Sur.) Hariot ssp. tomentosoides (van Goor) Silva, on the native snail, Lacuna vincta (Montagu), in the Gulf of Maine. The Veliger, No. 47:85-90.

      Chavanich S; Harris LG; Je JongGeel; Kang RaeSeon, 2006. Distribution pattern of the green alga Codium fragile (Suringar) Hariot, 1889 in its native range, Korea. Aquatic Invasions, 1(3):99-108. http://www.aquaticinvasions.ru/2006/AI_2006_1_3_Chavanich_etal.pdf

      Churchill AC; Moeller HW, 1972. Seasonal patterns of reproduction in New York populations of Codium fragile (Sur.) Hariot subsp. tomentosoides (Van Goor) Silva. Journal of Phycology, No. 8:147-152.

      Dromgoole FI, 1975. Occurrence of Codium fragile subspecies tomentosoides in New Zealand waters. New Zealand Journal of Marine and Freshwater Research, No. 9:257-264.

      Fletcher RL; Blunden G; Smith BE; Rogers DJ; Fish BC, 1989. Occurrence of a fouling, juvenile, stage of Codium fragile ssp. tomentosoides (Goor) Silva (Chlorophyceae, Codiales). Journal of Applied Phycology, No. 1:227-237.

      Fralick RA; Mathieson AC, 1972. Winter fragmentation of Codium fragile (Suringar) Hariot ssp. tomentosoides (van Goor) Silva (Chlorophyceae, Siphonales) in New England. Phycologia, No. 11:67-70.

      Fralick RA; Mathieson AC, 1973. Ecological studies of Codium fragile in New England, USA. Marine Biology, No. 19:127-132.

      Guiry MD; Guiry GM, 2007. AlgaeBase. Galway, Ireland: National University of Ireland. http://www.algaebase.org

      Hanisak MD, 1979. Growth patterns of Codium fragile ssp. tomentosoides in response to temperature, irradiance, salinity, and nitrogen source. Marine Biology, No. 50:319-332.

      Hanisak MD, 1979. Nitrogen limitation of Codium fragile ssp. tomentosoides as determined by tissue analysis. Marine Biology, No. 50:333-337.

      Harris LG; Jones AC, 2005. Temperature, herbivory and epibiont acquisition as factors controlling the distribution and ecological role of an invasive seaweed. Biological Invasions, 7(6):913-924. http://www.springerlink.com/content/l6871206466n4655/fulltext.pdf

      Harris LG; Mathieson AC, 2000. Marine Bioinvasions [ed. by Pederson J]. USA: Massachusetts Institute of Technology, MA, 46-56.

      Harris LG; Tyrrell MC, 2001. Changing community states in the Gulf of Maine: Synergism between invaders, overfishing, and climate change. Biological Invasions, No. 3:9-21.

      Hubbard CB; Garbary DJ, 2002. Morphological Variation of Codium fragile (Chlorophyta) in Eastern Canada. Botanica Marina, No. 45:476-485.

      Kang JW, 1966. On the geographical distribution of marine algae in Korea. Bulletin of Pusan Fisheries College, no. 7:1-125.

      Lee I; Kang J, 1986. A checklist of marine algae in Korea. The Korean Journal of Phycology, No. 1:311-325.

      Malinowski KC; Ramus J, 1973. Growth of the green alga Codium fragile in a Connecticut estuary. Journal of Phycology, No. 9:102-110.

      Mathieson AC; Dawes CJ; Harris LG; Hehre EJ, 2003. Expansion of the Asiatic green alga Codium fragile subsp. tomentosoides in the Gulf of Maine. Rhodora, 105(921):1-53.

      Moeller HW, 1969. PhD thesis. New Jersey, USA: Rutgers University.

      Neill PE; Alcalde O; Correa J, 2003. Proceedings of the XLVI Reunión Anual de la Sociedad de Biología de Chile, XVI Reunión Anual de la Sociedad de Ecología de Chile, Puyehue, Chile, R-56.

      Oh YS; Lee LP; Lee IK, 1987. A taxonomic study on the Genus Codium, Chlorophyta, in Cheju Island. The Korean Journal of Phycology, No. 1:61-72.

      Prince JS, 1988. Sexual reproduction in Codium fragile ssp. tomentosoides (Chlorophyceae) from the northeast coast of North America. Journal of Phycology, No. 24:112-114.

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      21/12/07 Original text by:

      Suchana Chavanich, Chulalongkorn University, Department of Marine Science, Faculty of Science, Bangkok 10330, Thailand

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