Invasion HistoryFirst Non-native North American Tidal Record: 1999
First Non-native West Coast Tidal Record:
First Non-native East/Gulf Coast Tidal Record: 1999
General Invasion History:
The origin of Didemnum perlucidum is unknown. It was first described in Guadeloupe in the Caribbean (Monniot 1983), but was subsequently found in Brazil, West Africa, the Gulf of Mexico, and the Indo-Pacific, including Hawaii, Guam, and the Pacific entrance to the Panama Canal (da Rocha and Kremer 2005). We consider this tunicate to be cryptogenic (status unknown) over much of its range, but there are several recent occurrences in harbors or on man-made structures where it is known to be introduced. These include the Gulf of Mexico, Florida, Guam, Hawaii, and the Pacific Coast of Panama (Godwin and Lambert 2000; Lambert 2002; Carman et al. 2010).
North American Invasion History:
Invasion History on the West Coast:
Invasion History on the East Coast:
In 1999, divers surveying an abandoned oil platform, converted to an artificial reef, off the coast of Texas found that it was incrusted with an unfamiliar colonial tunicate, which was identified by Gretchen Lambert as D. perlucidum (Culbertson and Harper 2001; Lambert 2002). Colonies of D. perlucidum were also tentatively identified on the SERC Invasions program fouling plates in Pensacola Bay (2002), Miami Harbor (2004), and the Indian River Lagoon (2005; 2010) (Ruiz et al. unpublished data). However, some of these colonies were young or in poor condition. D. perlucidum appears to be established in US waters, but its extent and abundance is unclear.
Invasion History on the Gulf Coast:
In 1999, Didemnum perlucidum was found on an abandoned oil platform/artificial reef off the coast of Texas (Culbertson and Harper 2001; Lambert 2002) and was tentatively identified by the Smithsonian Environmental Research Center’s Marine Invasions Lab in Pensacola Bay (in 2002) (Ruiz et al. unpublished data).
Invasion History in Hawaii:
Populations of Didemnum perlucidum, confined to harbors and artificial structures, were reported in 1999 from Ke'ehi Lagoon and Kaneohe Bay, Hawaii (Godwin and Lambert 2000; Coles et al., 2002).
Invasion History Elsewhere in the World:
Didemnum perlucidum was described from the undersides of buoys in a marina in Guadeloupe, in the West Indies by Francoise Monniot (1983). It was found in Sao Sebastiao Channel, southern Brazil in 1994. It has been collected from Pecem and Mucuripe Harbors, Ceara state, northern Brazil, (2009, Lotufo and Oliveira Filho 2010), from Isla Margarita, Venezuela (2009, da Rocha et al. 2010), and Pelican Cay, Belize (1992, Goodbody 2000). In the eastern Atlantic, it was collected in 1990 on Madeline Island, off Dakar, Senegal (Monniot and Monniot 1994).
In the Indo-Pacific, D. perlucidum has been collected in harbors in Zanzibar and Tanzania, Monniot and Monniot 1997), in the Maldives, in Indonesia and the Philippines (Monniot and Monniot 2001), and New Caledonia (Monniot and Monniot 1996). Among the Pacific islands, populations confined to harbors and artificial structures occur in Palau and Guam (Lambert 2002; Lambert 2003), Hawaii (1999, Ke'ehi Lagoon and Kaneohe Bay, Godwin and Lambert 2000; Coles et al., 2002a), and Papeete, Tahiti (Monniot et al. 1985). In the eastern Pacific, this tunicate was reported from natural rock wall communities (Witman and Smith 2003) and fouling plates (Lambert 2019) in the Galapagos Islands, and from Panama Bay, near the Pacific entrance to the Panama Canal (2008, Ruiz et al., unpublished data; Carman et al. 2010). A recent genetic study found a low level of genetic diversity, with only 3 mitochondrial COI haplotypes over the whole known range, and only 1 haplotype in Brazil and Australia. Notable new records in this survey were Veracruz, Mexico, in the Gulf of Mexico, Mazatlan, on the Mexican Pacific coast, and Kochi, Shikoku, Japan, where it was co-existing with D. vexillum (Dias et al. 2016).
Didemnum perlucidumis a colonial tunicate. Colonial tunicates are communities of individuals, called zooids, which share a protective cellulose layer called a tunic. Didemnum perlucidum contains spicules; small crystalline structures embedded in the tunic. Colonies vary greatly in appearance and the number of spicules they contain. The largest colonies measure 8 cm but are only 1-3 mm thick, and are usually found encrusting rock, wood, ropes, or other structures (Monniot 1983). Colonies are marbled in appearance—white and gray, yellow, or brown—with the darker color due to fecal pellets visible in the cloacal canals, where the tunic has only sparse spicules. According to Monniot (1983), some colonies completely lack spicules, but the colonies observed by da Rocha and Monniot (1995) in Brazil, always had them. Spicules occasionally reach 40µm in size, but usually do not exceed 20–30 µm (Monniot 1983; da Rocha and Monniot 1995). The zooids are arranged in clumps, leading to meandering dark lines on the tunic surface which are areas lacking in spicules between the clumps of zooids.
The zooids vary in appearance among colonies and the height of the colony is variable, but average around 1 mm (Monniot 1983; da Rocha and Monniot 1995). The oral siphon is short, large, and divided into six pointed lobes. The cloacal siphon varies in width, from 'modest' to exposing the whole branchial sac. There is an oval stomach followed by an annular post-stomach. The middle intestine is folded against the most posterior part of the abdomen. The single testis consists of a spherical body, surrounded by five to seven coils of the sperm duct. The ovary consists of one large oocyte and several smaller ones. The larvae are small (0.4-0.5 mm in diameter) in relation to the zooids, bearing three long adhesive papillae and four pairs of ampullae. The tail is long and forms a coil around the larva (Monniot 1983; da Rocha and Monniot 1995).
Potentially Misidentified Species
Probably native to Indo-Paciifc, broadly distributed. One disitnguishing feature is a gritty texture, due to embedded spicules.
Native to NW Pacific, widespread, mostly in col-water regions, but co-occurs with D. perlucidum in southern Japan.
Life History- A colonial tunicate consists of many zooids, bearing most or all of the organs of a solitary tunicate, but modified to varying degrees for colonial life. Colonial tunicates of the family Didemnidae have small zooids, completely embedded in an encrusting and thin tunic. Each zooid has an oral siphon and an atrial aperture which opens to a shared cloacal chamber. Water is pumped into the oral siphon, through finely meshed ciliated gills on the pharynx, where phytoplankton and detritus is filtered, and passed on mucus strings to the stomach and intestines. Excess waste is expelled in the outgoing atrial water (Van Name 1945; Barnes 1983).
Colonial tunicates reproduce both asexually by budding and sexually from fertilized eggs that develop into larvae. Buds can form from the body wall of the zooids. Colonies vary in size ranging from small clusters of zooids to huge spreading masses. The zooids are hermaphroditic, which means both eggs and sperm are released into the atrial chamber. Eggs may be self-fertilized or fertilized by sperm from nearby animals, but some species have a partial block to self-fertilization. Fertilized eggs are brooded within the tunic until they hatch into lecithotrophic (non-feeding, yolk-dependent) tadpole larvae. The larva has a muscular tail and a notochord, eyespots, and a set of adhesive papillae. The larvae are expelled upon hatching and swim briefly before settlement. Swimming periods are usually less than a day, but some larvae settle immediately after release or swim for longer periods if the water temperature is low. On settlement the tail is absorbed, the gill basket expands, and the tunicate begins to feed by filtering (Van Name 1945; Barnes 1983).
Didemnum perlucidum is known from tropical and subtropical climates in marine waters on rocks, coral reefs, shellfish culture operations, oil platforms, marinas, and docks (Culbertson and Harper, 2001; Lambert 2002; da Rocha et al. 2009; Sheehy et al. 2009). Large biomasses of colonial tunicates can filter large volumes of water, removing phytoplankton and other particles.
|General Habitat||Coarse Woody Debris||None|
|General Habitat||Marinas & Docks||None|
|General Habitat||Coral reef||None|
|General Habitat||Vessel Hull||None|
|General Habitat||Grass Bed||None|
|Salinity Range||Polyhaline||18-30 PSU|
|Salinity Range||Euhaline||30-40 PSU|
|Tidal Range||Low Intertidal||None|
Tolerances and Life History Parameters
|Broad Temperature Range||None||Warm temperate-Tropical|
|Broad Salinity Range||None||Polyhaline-Euhaline|
Shipping and Fisheries: Didemnum perlucidum is known to grow on boat hulls and on cultured mussels, such as Perna perna in Brazil. It overgrows mussels and could damage the bivalve industry; however, it was common but not dominant in mussel farms in southern Brazil (da Rocha et al. 2010).
Competition: On artificial substrates such as oil platforms, Didemnum perlucidum settles on or overgrows a wide range of organisms. On an abandoned oil platform in the Gulf of Mexico it appeared to settle on and overgrow sponges, corals, bryozoans, hydroids, and mollusks (Culbertson and Harper 2001). In fouling plate experiments in Sao Sebastiao, Brazil, it grew rapidly on bare plates and was able to overgrow barnacles and bivalves in about 70% of the cases and algae, bryozoans, and other colonial tunicates in about 65% of cases (Dias et al. 2008). In similar experiments in Santa Catharina, Brazil the barnacle Megabalanus coccopoma and the solitary tunicate Styela plicata survived overgrowth (Kremer et al. 2010) and it did not influence taxonomic richness or inhibit colonization by any species and appeared to have only weak, sporadic effects. These weak effects may have been due to the low abundance of D. perlucidum at this site (Kremer and Rocha 2011). This tunicate is known to grow on natural substrates in Guadeloupe and Brazil, but colonies are rare and smaller than those that occur on artificial structures (Monniot and Monniot 1985; Kremer et al. 2010). In the Swan River estuary, Western Australia, D. perlucidum has been spreading from artificial structures to seagrass beds (Halophila ovalis), reducing growth and photosynthesis, and decreasing habitat for the nonindigenous snail Batillaria ovalis, and probably for native species, as well (Simpson et al. 2016).
|CAR-I||Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida||Ecological Impact||Competition|
|On an abandoned oil platform in the Gulf of Mexico, Texas, Didemnum perlucidum appeared to settle on or overgrow 'sponges, corals, bryozoans, hydroids, and mollusks' (Culbertson and Harper 2001).|
|In fouling plate experiments in São Sebastião, Brazil, Didemnum perlucidum grew most rapidly on bare plates, and when other organisms settled, it grew at a much slower rate, but overgrew solitary organisms (barnacles, bivalves) in about 70% of the cases. However, encounters with algae, bryozoans, or colonial ascidians resulted in stand-offs in about 65% of cases (Dias et al. 2008). In similar experiments in Santa Catarina state, overgrown species, such as the barnacle Megabalanus coccopoma and the solitary ascidian Styela plicata survived overgrowth (Kremer et al. 2010). Didemnum perlucidum did not influence taxonomic richness or inhibit colonization by any species, and appeared to have only weak, sporadic effects (Kremer et al. 2010). These weak effects may have been due to low abundance at this site (Kremer and Rocha 2011). In a 3-species experimnent, D. perlucidum dominates over Botrylloides nigrer and Schizoporella errata when preators were excluded, during the summer (Oricchio and Dais 2020).|
|Didemnum perlucidum was common but not dominant in mussel farms in southern Brazil. It does overgrow mussels, so could 'be damaging to the bivalve industry' (da Rocha et al. 2010). In later studies, D. pelucidum was found to affect the growth of cultuered mussels, delying or preventing their growth to maretable size. Monthly cleaning of the mussels and the culture 'socks' improved the growth of the mussels, but increased labor costs, (Lins and Rocha 2020).|
|AUS-IV||None||Ecological Impact||Habitat Change|
|Didemnum perlucidum has been overgrowing blades of the seagrass Halophila ovalis in the Swan River estuary, Western Australia, decreasing seagrass cover. Density of the nonindigenous snail Batillaria australis has decreased in areas of grass beds colonized by D. perlucidum (Simpson et al. 2016).|
|Didemnum perlucidum has been overgrowing blades of the seagrass Halophila ovalis in the Swan River estuary, Western Australia, decreasing the plant's rate of photosynthesis and growth. The tunicate has been spreading outward from areas of human infrastucture, such as docks and buoys (Simpson et al. 2016).|
|Predator exclusion and natural (upwelling) eutrophication allowed D. cf. perlucidum to overgrow corals, polychaetes, and barnacles (Roth et al. 2017).|
|SEP-H||None||Ecological Impact||Habitat Change|
|Rapid overgrowth of fouling communities by D. cf. perlucidum can potentially affecting the composition of the coral reef community (Roth et al. 2017).|
Regional Distribution Map
|Bioregion||Region Name||Year||Invasion Status||Population Status|
|CAR-I||Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida||1999||Def||Estab|
|PAN_PAC||Panama Pacific Coast||2008||Def||Estab|
|PAN_CAR||Panama Caribbean Coast||2008||Def||Estab|
|6828||Culbertson and Harper 2000||1999||1999-01-01||12 miles east of Stetson Bank||Def||28.2500||-94.4667|
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