Tubastraea coccinea

Invasion History

First Non-native North American Tidal Record: 1977
First Non-native West Coast Tidal Record:
First Non-native East/Gulf Coast Tidal Record: 1977

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General Invasion History:

Tubastraea coccinea has a broad Indo-Pacific range, from the Red Sea and Madagascar, to the west coast of the tropical Americas from the Gulf of California to Peru. Transport by ships is possible over some of this range, but cannot be documented (Creed et al. 2016). It was described from the Red Sea in 1829, and was collected in Acapulco, Mexico before 1866 (Verrill 1866, cited by Cruz-Pinon and Reyes Bonilla 1999). Before 1953, it was introduced to the Caribbean, and has spread north to oil platforms off Texas and Louisiana, and to the coast of south Florida (Cairns 2000; Fenner 2001; Fenner and Banks 2004; Creed et al. 2016). By 1990, it was discovered in Brazil, where it has been spreading (Figueira de Paula and Creed 2005; Creed 2006; Creed et al. 2016). Hull fouling has probably been the major vector of transport between distant ocean regions, but the larvae can remain competent for 100 days, permitting regional dispersal by currents (Fenner 2001). Web searches indicate that aquarists also cultivate and maintain these attractive corals, although, since they lack symbiotic algae, they require regular feeding with brine shrimp or artificial foods. Disposal by aquarists is a potential vector for future invasions (Shearer 2011; Creed et al. 2016).

North American Invasion History:

Invasion History on the East Coast:

On the East Coast of the US, T. coccinea was first found in 1999, on a shipwreck off Key Largo, Florida (Fenner and Banks 2004). It now inhabits many shipwrecks, and several limestone mitigation reefs from the northern Florida Keys to the Palm Beach inlet. It can be very abundant on artificial substrate, but has not yet been found on natural surfaces (Shearer 2011). In 2015, it was found growing on a buoy in the Gray's Reef National Marine Sanctuary, off Georgia (Creed et al. 2017).  A single specimen was found on a buoy at Grays Reef National Marine Sanctuary off Georgia (S. Fangman, cited by Creed et al. 2017).

Invasion History on the Gulf Coast:

In 1977, T. coccinea was found in the Gulf of Mexico on oil drilling platforms 50–100 km off Tuxpan, in Veracruz, Mexico. In 1991, it was found on oil platforms off Texas (Fenner 2001) and in 1994, it was found at a platform near the Mississippi Canyon, off Louisiana. By 2000, it had been found at 11 offshore platforms. In 2002, T. coccinea was observed on the East Flower Garden Bank, a deep-water coral reef and National Marine Sanctuary in the Northwestern Gulf of Mexico (Fenner and Banks 2004). In 2010, this coral was observed growing on natural substrates in the West Flower Garden Banks (Shearer 2011).Modeling suggests that surface pH, calclite, and benthic current velocity, iron concentration, and dissolved oxygen infleuenced T. coccinea abundance, predicting greatest abundance on the coasts of the western Gulf of Mexico from Louisiana to Texas, an area with a high concentration of oil platforms (Derouen et al. 2020).

Invasion History Elsewhere in the World:

Tubastraea coccinea reached Jamaica by 1955 (Cairns 2000) and by the 1970s was found on the southern coast of the Caribbean. By 1999 it had reached Belize and Cozumne (Fenner 2001). In Brazil, T. coccinea was first found on offshore oil rigs over the Campos Basin in Brazil, north of Rio de Janeiro (1990, Figuera de Paula and Creed 2004). In 2008, this coral was found 130 km further south, in the Buzios and Vittoria Islands (Mantelatto et al. 2011). Also in 2008, T. coccinea was found on shipwrecks, marinas, and coral reefs near Salvador, Bahia, about 1200 km north (Sampaio et al. 2012). On the coast of Brazil, this coral now ranges over 3500 km. Some genotypes are widespread, but there is evidence for multiple introductions, particularly on offshore oil platforms arriving from the Indo-Pacific (Capel et al. 2019). Tubastraea concinnea has been identified and is spreading in the Canary Islands, wiht a first record in 2015 (Brito et al. 2017; Lopez et al. 2019). It has been reported from West Africa, from the Cape Verde Islands and the Gulf of Guinea, but the identification is questionable (Cairns 2001). The Cape Verde-West African form has been recognized as a new genus and species, Atlantia caboverdiensis (Capel et al. 2020).

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Description

Tubastraea coccinea is an azooxanthellate coral (lacking symbiotic algae), which grows in colonies consisting of clumps of calcareous cups, projecting from a spongy calcareous base. The corallite (calcareous wall around a single polyp) is cylindrical, up to 11 mm in diameter, and may be flush with the colony skeleton (coenosteum) or project up to 4 cm above it. The synapticulotheca (wall of the corallite) is porous and lacks a covering layer (epitheca). The corallite skeleton is white, with poorly defined ribs. The interior of the corallite is partly divided by septa, projecting into the body cavity; there are four kinds of septa, of varying width (S1>S2 >S3>S4), which are arranged in cycles around the opening of the corallite. There are 48 septa. The septa do not protrude on the exterior of the corallite. There is a prominent columella, a projection of the skeleton in the center of the corallite. The body of the polyp itself is deep red or orange, while the tentacles are yellow to bright orange (description from: Kaplan 1988; Cairns 2000; Figueira de Paula and Creed 2004).

The colonies of T. coccinea can vary greatly in arrangement, probably as a result of their environment. They can be cerioid, composed of corallites united directly to one another by fused walls, with corallites that are juxtaposed; plocoid, composed of more or less cylindrical corallites having distinct walls, and the coenosteum separates corallites within a colony; or phaceloid, composed of laterally free corallites forming tufts, and the corallites are separated by void space. In shallow waters, subject to waves and currents, colonies are usually cerioid to plocoid and tightly arranged, but in deeper calmer waters they are more likely to be phaceloid and loosely arranged, with widely projecting corallites. Colonies tend to form spherical or mound-shaped clumps, up to 14 cm in diameter, and strongly attached to the substrate (description from: Cairns 2000; Figueira de Paula and Creed 2004).

Two morphotypes of T. coccinea were observed on an oil platform in the Gulf of Mexico, a dark-orange red form with corallites not projecting far above the colony surface (plocoid, 'low' morphotype), and a yellow form with corallites projecting and loosely arranged (phaceloid, 'high' morphotype). Both forms (examined by Stephen Carins) had skeletal morphology corresponding to T. coccinea, but they show differences in distribution, substrate preference, reproductive timing, and could represent cryptic species (Shearer 2011).

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Taxonomy

Ecology

General:

Planulae in Brazil settled equally on plates made with wood, granite, concrete, steel and ceramic tiles (Creed and Figueira de Paula 2007). In Brazil and the Caribbean, this coral is common on natural substrates, though preferring artificial ones (Mangelli et al. 2012). However, in the Gulf of Mexico most occurrences have been on shipwrecks, oil platforms, and dock floats (Cairns 2000; Fenner 2001; Fenner and Banks 2004; Creed and de Paula 2007; Shearer 2011), but occurrences are also known from natural substrates on the Flower Garden Banks (Fenner and Banks 2004). In experiments, T. coccinea survived exposure to temperatures as low as 15 °C for 96 hours, but died in 24-hour exposure at 12.5 °C. Its distribution on Cabo de Frio Island, Brazil, was strongly affected by an adjacent cool-water upwelling zone, and was rare or absent where temperatures dropped below 15 °C (Batisita et al. 2016). Tubastraea coccinea lacks zooxanthellae (symbiotic algae), and so is not dependent on photosynthesis. Because they are not limited by light, they can occur at depths down to 60m (Shearer 2011). They feed on zooplankton captured with their tentacles.

Food:

Zooplankton

Consumers:

Competitors:

Trophic Status:

Primary Producer

PrimProd

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Habitats

General Habitat	Marinas & Docks	None
General Habitat	Vessel Hull	None
General Habitat	Rocky	None
General Habitat	Coral reef	None
Salinity Range	Polyhaline	18-30 PSU
Salinity Range	Euhaline	30-40 PSU
Tidal Range	Subtidal	None
Vertical Habitat	Epibenthic	None

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Life History

Tubastraea coccinea is a coral which grows in small colonies (up to 14 cm in diameter), but does not form reefs. Polyps reproduce asexually by budding from the base or from the oral disk to produce new polyps, which secrete their own calcareous skeletons (Barnes 1983; Harrison 2011). Polyps of this coral are hermaphroditic, and release sperm into the water, but brood their eggs, which develop into planula larvae. This coral can also produce planulae asexually (Ayres & Resing 1986; Harrison 2011). Local populations include a high proportion of clones, indicating that asexual reproduction contributes to the rapid spread of this coral (Capel et al. 2017). The planulae are quite large (3–5 mm long) and can be visible during mass-spawning events (Paz-Garcia et al. 2007). They can remain competent to settle for up to 18 days, so have the capability for long-distance dispersal by currents (Fenner 2001; Figueira de Paula et al. 2014). Fragments of T. coccinea have a remarkable ability to regenerate into living, whole polyps, which has implications for control efforts (Luz et al. 2018).

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Tolerances and Life History Parameters

Maximum Depth (m)	78	Gulf of Mexico, off Louisiana (Sammarco et al. 2013)
Minimum Temperature (ºC)	15	Experiment, 96 h exposure, no mortality. 100% mortality in 24 h occurred at 12.5 C (Batista et al. 2016).
Minimum Duration	1	Time to settlement (Glynn et al. 2008)
Maximum Duration	18	Time to settlement is usually 3-5 days(Glynn et al. 2008), but larvae in aquaria can remain competent for up to 18 days (Figueira de Paula et al. 2013).
Broad Temperature Range	None	Subtropical-Tropical

General Impacts

Tubastraea coccinea has been spreading rapidly in the tropical Western Atlantic, and in the subtropical Gulf of Mexico. In US waters, it has largely been confined to artificial substrates such as oil platforms, shipwrecks, and artificial reefs, but its spread to coral reefs is a concern, particularly in the Flower Garden Banks National Marine Sanctuary (Fenner 2001; Fenner and Banks 2004; Sammarco et al. 2004). A number of impacts have been studied in Brazilian waters, including: competition and overgrowth of native corals (Creed 2006; Silva et al. 2011); how settlement affects food webs by replacing hermatypic corals (containing symbiotic algae, relying partly on photosynthesis for nutrition) with corals which are exclusively carnivorous (Silva et al. 2011); and the inhibition of feeding by coral reef fishes (Lages et al. 2011). Modeling of the environmental requirements of T. coccinea suggest that it will greatly expand its range in Brazil, and has the potential to compete with the native coral Mussismilia hispida (Riul et al. 2013). Wrapping the corals with plastic or raffia has been found to be an effective means of controlling isolated infestations of T. coccinea and T. tagusensis (Mantelatto et al. 2015).

In laboratory experiments, T. coccinea extruded mesenteric filaments, and produced extracellular digestive responses to the native Florida anthozoans Ricordea florida (Cnidaria, Corallimorpharia), and the anemone Epicystis crucifer. Tubastraea coccinea caused tissue necrosis in the coral and anemone, and showed general patterns of aggressive behavior. However, no immune response or aggression was observed with another introduced coral, T. micranthus (Hennessey and Sammarco 2014).

Tubastraea coccinea is beautiful, and frequently photographed by divers in its native and introduced ranges. Although it lacks symbiotic algae, and requires daily feeding by hand, it is cultivated by aquarists (Shearer 2011). However, the aquarium trade provides another potential vector for this and other invasive coral species.

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Regional Impacts

SA-II	None		Ecological Impact	Food/Prey
Extracts of Tubastraea coccinea inhibit feeding by coral reef fishes (on tuna embedded in gel plates) (Lages et al. 2010). Field studies indicate that T. coccinea and T. tagusensis have little or no predation compared to native corals and sponges (Moreira and Creed 2012). Observations with remote cameras confirm that high densities of Tubastraea coccinea and T. tagusensis decreased feeding rates of territorial and mobile invertebrate-feeding fishes (Miranda et al. 2018).
SA-II	None		Ecological Impact	Habitat Change
Extracts of Tubastraea coccinea inhibit settLement by algae on plates, but favored one species of hydroid (Lages et al. 2010). Tubastraea coccinea and the introduced T. taguensis caused necrosis when in contact with the native coral Mussismilia hispida, creating dead areas which were overgrown by sponges and sometimes by Tubastrea recruits in Ilha Grande Bay, Brazil (Creed 2006). Rocky reefs heavuly colonized by Tubastraea cocinea and T. tagusensis had greatly reduced abundances of small mobile invertebrates (Silva et al. 2019). In experiments,  The invasion of T. coccinea and T. tagusensis has created habitat for the invasive bivalve Leiosolonus aristatus (Vinagre et al. 2017; Capel et al. 2020).
SA-II	None		Ecological Impact	Competition
Tubastraea coccinea and the introduced T. taguensis caused necrosis when in contact with the native coral Mussismilia hispida, creating dead areas which were overgrown by sponges and sometimes by Tubastrea recruits in Ilha Grande Bay, Brazil (Creed 2006). A later survey documented the continued spread of T. coccinea and T. tagusensis into the Tamoios Ecological Station Marine Protected Area in Ilha Grande Bay (Silva et al. 2011). Densities of T. coccinea in random quadrats increased in Ilha Grande Bay over a 2-year period, although it remained less abundant than native corals, ranking 16th among sessile benthic taxa (Lages et al. 2011). Tubastraea coccinea is now spreading in mussel (Perna perna) beds in Ilha Grande Bay (Mantelatto and Creed 2015). On Arvoredo Island, Brazil, T. coccinea dominates caves, crevices and walls, while the native Palythoa caribaeorum covers open areas. Competition here, at its southern limit, is limited by tmeprature stress (Almeida Saa et al. 2019; Guilehm et al. 2020)
SA-II	None		Ecological Impact	Trophic Cascade
Beacause T. coccinea and T. taguensis are hermatypic (lacking photosynthetic algae, zooxanthellae) and are replacing a native hermatypic coral (native coral Mussismilia hispida), their continued invasion threatens to lower the productivity of the reef (Silva et al. 2011). Increasing cover of Tubastrea spp. resulted in decreaed algal cover, and a decreasing abundance of roving herbivorous fishes (Miranda et al. 2018).
SA-II	None		Economic Impact	Fisheries
Tubastraea coccinea and T. tagusensis are now spreading in mussel (Perna perna) beds in Ilha Grande Bay. These corals are a possible threat to local mussel fisheires(Mantelatto and Creed 2015).
SA-II	None		Economic Impact	Ecosystem Services
In Brazil, invasions of Tubastraea spp. are perceived as negatively affecting the biodiversity, visual quality, and fisheries of rocky reefs and coral reefs of coastal waters (Dutra et al. 2024).  Consequently, there's been efforts to monitor its distribution and efforts at controlling it through local eradication (Creed et al. 2017; Dutra et al. 2023).  Methods have included the use of fresh or low-salinity water (Moreira et al. 2014(, wrapping colonies (Mantelatto et al. 2015), or use of sodium hypochlorite (Altvater et al. 2017).

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