Didemnum perlucidum

Invasion History

First Non-native Panama (Pacific) Tidal Record: 2008
First Non-native Panama (Caribbean) Tidal Record: 2009

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Panama Invasion History:

Invasion History in Panama:

Although Didemnum perlucidum was described from the Caribbean, its rapid spread in the Western Atlantic, its association with artificial structures, and its wide distribution in the Pacific, suggest that it is not native to the Caribbean (Lambert 2002; da Rocha and Kremer 2005; Dias et al. 2013). In 2008, Didemnum perlucidum was collected in Colon, Panama, at the Caribbean end and in Panama Bay, at the Pacific end (Carman et al. 2011). 

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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2019) in the Galapagos Islands, and from Panama Bay, near the Pacific entrance to the Panama Canal (2008, Ruiz et al., unpublished data; Carman et al. 2010).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).

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Description

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). 

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Taxonomy

Ecology

General:

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.

Food:

Phytoplankton; Detritus

Trophic Status:

Suspension Feeder

SusFed

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Habitats

General Habitat	Coarse Woody Debris	None
General Habitat	Marinas & Docks	None
General Habitat	Coral reef	None
General Habitat	Rocky	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	Subtidal	None
Tidal Range	Low Intertidal	None
Vertical Habitat	Epibenthic	None

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Life History

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Tolerances and Life History Parameters

Broad Temperature Range	None	Warm temperate-Tropical
Broad Salinity Range	None	Polyhaline-Euhaline

General Impacts

Economic Impacts

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).

Ecological Impacts

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).

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