Known introduction to Australia

General Information

(M.Vahl) C. Agardh, 1817

Please use the following citation for this material
NIMPIS 2013, Caulerpa taxifolia general information, National Introduced Marine Pest Information System, viewed 26 May 2013 <http://www.marinepests.gov.au/nimpis>.

Habitat

Found on a wide variety of substrates from rock, sand and mud to dead seagrasses. It occurs most frequently found in depths of 2.5-15 m but has been found to a depth of 100 m in the Mediterranean. The invasive aquarium strain is able to occupy up to 100% of the available substratum. Native Australian populations in tropical waters are found on rocky reefs and seagrass meadows in sheltered or moderately wave-exposed areas in both polluted and pristine waters.

Habitat classification

For the purposes of NIMPIS, habitats have been divided into four categories: Hard, soft, water and organism. The habitat types assigned to these categories reflect the variety of substrata available for organisms to colonise. Habitat types listed for this species are those that have been recorded in the literature.

Hard

This category contains both natural and artificial habitats that are solid, fixed or permanent substrata. Species can reside on (e.g. attached externally) or within (e.g. boring into) the habitat type.

• Bedrock
  

Soft

This category contains habitats that are not fixed or permanent, and are able to be affected by water movement. Species can reside on (e.g. living on the sediment-water interface) or within (e.g. burrowing into) the habitat type.

• Sand-fine
  
• Sand-medium
  

Organism

This category is used for species that reside on or within another organism.

• ism-Organism
  

Tidal Range

• Sub-tidal
  

Habitat survival parameters

	Minimum	Maximum
Adult temperature	7.0 °C The following temperatures have been recorded for survival and growth of the aquarium strain in the Mediterranean: survives as low as 7 °C (Komattsu et al 1994 in Komatsu et al. 1997, Boudouresque et al. 1995); no growth 10-12.5 °C, grows once temperatures reach 15 °C, when the MorphologyLength of stolons begins to increase. Blade growth begins at: 17.5 °C (Komatsu et al. 1997, Boudouresque et al. 1995); 18 °C, ambient temperature (Augier et al. 1982); 12 °C, ambient winter temperature (Chishom et al. 2000); 11 °C , ambient winter temperatures Monaco (Meinesz and Hesse 1991); 12 °C ambient winter temperatures Mediterranean (Chishom et al. 2000). 11 °C ambient winter temperatures Monaco (Meinesz and Hesse 1991). The lethal temperature of 9 °C has been recorded in the laboratory for cuttings cultured under controlled temperature and light (Komatsu et al., 1997).	32.5 °C The following temperatures have been recorded for survival and growth of the aquarium strain in the Mediterranean: optimum photosynthesis 20-30 °C (Komatsu et al 1994 in Komatsu et al., 1997); laboratory, cuttings cultured under controlled light at 31.5 °C (Komatsu et al 1994 in Komatsu et al., 1997); laboratory at temperatures of 17, 21, 25 °C and 38psu salinity, northern Mediterranean, May-October (Thibaut et al 2001). The lethal temperature of 32.5 °C has been recorded in the laboratory for cuttings cultured under controlled temperature and light (Komatsu et al., 1997).
Adult salinity	15.0 ppt In the laboratory, high mortality was found at salinity <15 ppt (Theil et al. 2007). However, level of mortality at 15 ppt was dependent on water temperature and Ca2+ concentration in water (the lower the temperature below 20 °C and the lower the Ca2+ concentration, the higher the mortality) with no growth evident in surviving plants. Exposure at 10 ppt for 180 min was fatal to all parts of the alga, while highest growth rates across temperatures up to 30 °C occurred at 25 ppt (Theil et al. 2007).	40.0 ppt In the laboratory, salinity levels of 38-40 ppt were survived without ill-effects (Williams and Schroeder 2004).
Dissolved oxygen	N/a	N/a

Reproduction and growth

Native populations of C. taxifolia are known to reproduce sexually, but the aquarium strain appears to be an all-male clone that only produces male gametes. In the Mediterranean, the aquarium strain spreads vegetatively by growth of the stolons or by regeneration from broken-off fragments as small as 1 square centimetre in size.

Minimum reproductive temperature	25.0 °C Zuljevic and Antolic(2000) (males only).
Maximum reproductive temperature	N/a
Cues	Gamete release occurred in the Adriatic Sea (Zuljevic & Antolic 2000)- all gametes were male and released within 10 minutes, about half an hour before sunrise (on overcast days - release was 20 mins before sunrise).
Strategy	Monoecious. In Mediterranean reproduction is vegetative by fragmentation (Komatsu et al., 1997). In native populations, male and females gametes fuse forming a zygote which grows through two little known stages prior to becoming the adult (Meinesz 1999).
Season	Sexual reproduction has been observed in central Queensland (pers. comm. Ian Price). Gametes released in June -September in Adriatic Sea (Zuljevic & Antolic 2000).

Life cycle

Age to maturity

Please use the following citation for this material
NIMPIS 2013, Caulerpa taxifolia reproduction and habitat, National Introduced Marine Pest Information System, viewed 26 May 2013 <http://www.marinepests.gov.au/nimpis>.

Feeding Preferences

Trophic status: primary producer

C. taxifolia produces its own food through photosynthesis.

Food

vegetative	C. taxifolia produces its own food through photosynthesis.

Competitors

Stage: vegetative	C. taxifolia competes with the seagrasses Posidonia oceanica and Cymodocea nodosa in Mediterranean ecosystems (Relini et al. 1998, Boudouresque et al. 1995).

Predators

In the Mediterranean, the C. taxifolia aquarium strain is eaten by the sea slugs Oxynoe olivacea, Lobiger serradifalci and Elysia subornata to some extent. Caulerpenyne, a toxin produced by C. taxifolia, appears to deter other grazing species.

Please use the following citation for this material
NIMPIS 2013, Caulerpa taxifolia feeding and predators, National Introduced Marine Pest Information System, viewed 26 May 2013 <http://www.marinepests.gov.au/nimpis>.

Impacts

In the Mediterranean Sea, the C. taxifolia aquarium strain is extremely invasive and can smother other algal species, seagrasses and sessile invertebrate communities. Sparse beds of other algae or seagrasses are particularly susceptible to invasion. Aquarium caulerpa out-competes other plants through vigorous stolon growth. Its large monospecific meadows have vastly reduced native species diversity and fish habitat, usually leading to reduction of catch rates of commercially valuable fish species.

Vectors

Descriptions of the vector types that are relevant to this species are displayed below.

Fisheries and Aquaculture

This class encompasses vectors associated with fisheries and aquaculture activities and trade. An example of a vector from this class is Fisheries intentional, which would incorporate, for example, the introduction of the Pacific oyster Crassostrea gigas for aquaculture purposes. Other vectors included in this class are: Discarded bait, Fisheries accidental (not mollusc), mollusc accidental, mollusc intentional, packing material and scientific escape.

Fisheries - accidental (not mollusc)	The accidental translocation of species through aquaculture and fisheries activities. This vector includes the accidental release of live fish, crustaceans and molluscs (other than oysters) imported for human consumption, This vector also includes the accidental translocation of species attached to aquaculture gear (floats, cages, etc).
Fisheries - intentional (not mollusc)	The deliberate translocation of fish, crustaceans or molluscs (not oysters) to establish or support a new fishery. For example many aquaculture operations use species that are not native, which involves introducing species from elsewhere in the world.
Oyster accidental	The introduction of organisms associated with molluscs and their deliberate translocation. Species of molluscs are highly valued aquaculture species and have been translocated across the globe for farming. Many species live cryptically on mollusc shells and have subsequently been introduced along with molluscs into new localities. This was more common historically when shells were not cleaned of other species.
Oyster intentional	This vector describes the deliberate introduction of molluscs. Species of molluscs are highly valued aquaculture species and have been translocated to many locations around the world to establish aquaculture industries and also to restock areas where native stocks have been lost through either disease or overfishing.
Discarded bait	The release of unused, unsuccessful or excess bait species (and associated organisms) from commercial fishing operations, sport and recreational fishing activities. It is possible that two species of prawn that are introduced into San Francisco Bay, were originally imported for sale as bait or human food. The dumping of organisms from vessels or release from shore may have led to the establishment in the wild.

Natural dispersal

Natural dispersal is a mechanism for the range expansion of a species through natural processes such as the movement of larvae or adults to a new location. As an example, through passive movement in water currents; or active movement (migration) in response to changes in environmental conditions such as salinity changes or water flow dynamics. Natural dispersal also allows for the successful settlement of recruits in a new location. The only vector associated with this class is: Natural dispersal

Ornamental

This class encompasses vectors associated with ornamental species in the aquarium trade or horticulture and landscaping industries. A vector from this class is Aquatic plant shipments. There are a large number of aquatic plants that have been introduced for decorative purposes in aquaria ponds (aquatic) or gardens (terrestrial) via these shipments. Other vectors that could be included in this class include: Garden escape, Pet release

Vessels

This class encompasses vectors associated with maritime transport and shipping activities. Vessels includes; commercial ships (e.g. tankers, container ships, ferries, barges), fishing vessels, recreational vessels, passenger vessels, drilling platforms and research vessels. An example of a vector from this class is ballast water,which has been found to transport up to 10 000 different species at any one time. Other vectors associated with this class include: dry ballast, biofouling community

Biofouling	Fouling communities are typically composed of encrusting or sessile species, however they can include mobile species. This vector can introduce species through a variety of means. Three examples are: (1) The spawning of a fouling species on a vessel in port and its successful settlement and establishment of a reproductive population; (2) The dislodgement of fouling species from a vessel in port through abrasion with wharf structures, ropes, etc., or through in water vessel hull cleaning (banned in Australia) or through high vessel speeds, etc.; and (3) The sinking of fouled vessels either deliberately or accidentally can introduce new species to a location. There are a variety of vectors capable of having a fouling community. Characteristics of a fouling community found on wooden boat hulls include: having a wood boring habit; a benthic sessile or encrusting stage; and mobile adults or larval stages. Fouling communities found within sea chests, anchor wells etc. often are mobile crevice occupying species or known obligate associate of fouling species and can escape into new locations.
Ballast water	The release of species in ballast water discharged from vessels. Various types and life stages of species can be transported in ballast water, including plankton, crustaceans, fish, larvae, eggs or cysts. Ballast water is used in commercial vessels to stabilise the vessel and is uploaded or discharged depending on the amount of cargo onboard. Ballast water as a vector also includes sediments that accumulate in the bottom of ballast tanks. Species that are able to survive within these sediments include those that have a resistant stage or resting cyst (eg. dinoflagellates) as well as adult stages of benthic organisms.
Dry ballast	The accidental release of species with solid ballast. Though solid ballast has predominantly been replaced by ballast water, it historically was used in vessels to stabilise the ship during transit. Dry ballast included rocks, sand, wood and other substrata collected from the foreshore and hence many intertidal species were also unintentionally included. When no longer required, this dry ballast was disposed of, usually overboard or onto the foreshore for subsequent use, releasing organisms to a new environment.
Fisheries - accidental (not mollusc)	The accidental translocation of species through aquaculture and fisheries activities. This vector includes the accidental release of live fish, crustaceans and molluscs (other than oysters) imported for human consumption, This vector also includes the accidental translocation of species attached to aquaculture gear (floats, cages, etc).
Packing material	The accidental release of species associated with seaweed (and other packing materials) for bait and fishery products. These packaging materials are often disposed of at sea by fishers, which can release organisms into the marine environment.
Fisheries - accidental (products)	The accidental translocation of species through aquaculture and fisheries activities. This vector includes the accidental release of live fish, crustaceans and molluscs (other than oysters) imported for human consumption, This vector also includes the accidental translocation of species attached to aquaculture gear (floats, cages, etc).

Please use the following citation for this material
NIMPIS 2013, Caulerpa taxifolia impacts and vectors, National Introduced Marine Pest Information System, viewed 26 May 2013 <http://www.marinepests.gov.au/nimpis>.

Additional Information

General Notes

vegetative

New stolons form in April-July (France), sexual and vegetative reproduction; toxic secondary metabolites form an efficient strategy against grazers; the major metabolite, caulerpenyne, is most active on sea urchin eggs. Habitat: All substrate types from rock, sand to mud, to depths of 3-35 m, but has been recorded to 90 m; on reef flats in sheltered or moderately wave-exposed areas. Dispersing of cuttings (vegetative reproduction) may occur naturally in the immediate area of a colonised site or artificially at greater distances by means of pleasure boat anchors or fishing nets (Komatus et al., 1997). Stolon growth of Mediterranean strain strongly correlated with increasing temperature. Vital temperature range 10-32.5 °C, no growth 10-12.5 °C, new stolon and frond development 15-17.5 °C. Most favourable range of light intensity 88-338 umol m-1s-1 (14 h light: 10 h dark) (see Komatsu et al., 1997). Morphological, ecological and physiological studies demonstrated that Mediterranean C. taxifolia differs from known tropical populations, exhibiting larger size, vigorous growth and resistance to low temperatures (Jousson et al., 1998). Found in sand on reef flats in sheltered or moderately wave-exposed areas to 15m deep (Scullion Littler et al., 1989). Caulerpa taxifolia and other species of the genus Caulerpa are known to show highly variable morphologies depending on environmental conditions such as light, nutrients and seasons (Chisholm et al. 1995). Caulerpa spp. Are known to reproduce sexually by fusion of male and female gametes and asexually by fragmentation. Zulevic & Antolic (2000) documented gamete formation and release for a Mediterranean population of C. taxifolia in Croatia - the gametes were all male, confirming the hypothesis that the introduced populations may be an all-male clone. The growth of C. taxifolia is pseudoperennial - intermediate between annual and perennial. Rhizome apical growth is indefinite, however no single part of the alga persists physically for more than one year (Meinesz et al. 1995) Studies conducted on native and invasive populations in Australia showed invasive populations to have higher levels of fronds arising from asexual fragmentation (Wright 2005).

Identification Notes

vegetative

Green algae; stolon diameter 1.5-1.8 mm, length <2.8 m, fronds compressed laterally, slightly constricted at base, upwardly curved and gradually tapering at apex, do not overlap, frond diameter 6-8 mm, rhizoid pillars spaced 1.6-4.3 cm apart, frond length from 3-15 cm in shallower lighter conditions to 40-60cm in deeper situations with maximum ramifications in spring (France), small lateral branchlets which are constricted at point of attachment but do not taper into midrib. C. taxifolia is distinguished from other Australian Caulerpa species by the facts that branchlets that are constricted at the base (where they attach to the midrib of each frond) and that they are opposite in their attachmentto the midrib (as opposed to alternating). Thirdly the branchlets curve upwardsand taper at the apex.

Similar Species

vegetative

Caulerpa mexicana, C. prolifera, C. racemosa. C. mexicana has been suggested as an ecad (ecological morph) of C. taxifolia- a variation of the species adapted to its surrounding environmental conditions (Chisholm et al. 1995). Caulerpa veravalensis, endemic to theArabian Sea, resembles C. taxifolia but differs in the shape of the stolon and the base of the assimilators (C. veravalensis is bulbous at the base, C. taxifolia is cylindrical at the base) (see Shameel 1978).

Please use the following citation for this material
NIMPIS 2013, Caulerpa taxifolia additional information, National Introduced Marine Pest Information System, viewed 26 May 2013 <http://www.marinepests.gov.au/nimpis>.

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