Carijoa riisei

Invasion History

First Non-native North American Tidal Record: 1869
First Non-native West Coast Tidal Record:
First Non-native East/Gulf Coast Tidal Record: 1869

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General Invasion History:

Carijoa riisei is a soft coral widely distributed in tropical and subtropical marine waters around the world. It was described from the Virgin Islands in 1860 and subsequently found in the Atlantic from South Carolina to Brazil (Deichmann 1936; DeVictor and Morton 2010; U.S. National Museum of Natural History 2014). In 1966, it was found in Oahu, Hawaii and by 2000, became established throughout the Hawaiian Islands, from shallow bays to depths of 110 m (Kahng and Grigg 2005; Carlton and Eldredge 2009). Carijoa riisei was presumed to be an invader of Caribbean origin, but a genetic study of worldwide populations found that genetic diversity of this coral was greatest in the Indo-West Pacific, less in the Hawaiian Islands, and least in the Western Atlantic (Concepcion et al. 2005). Consequently, C. riisei appears to be native to the Indo-Pacific, introduced to the Hawaiian Islands, and a very early introduction to the Western Atlantic. Based on the specimens examined by Concepcion et al. (2010), and museum specimens, the native range of C. riisei includes the Indian and western Pacific Oceans from the Gulf of Oman and Okinawa, south to Mozambique and South Australia, and west to Fiji and Tonga (Concepcion et al. 2010; U.S. National Museum of Natural History 2014).

North American Invasion History:

Invasion History on the East Coast:

Carijoa riisei was probably a very early introduction to the East Coast, since it was described from the Virgin Islands in 1860 (Deichmann 1936) and found in the Gulf of Mexico on the Dry Tortugas (in the Florida Keys) in 1869 (YPM IZ 002392 CN, Yale Peabody Museum 2014). The first East Coast record is from Biscayne Bay, Florida in 1947 (USNM 44065, U.S. National Museum of Natural History 2014). Other Florida records include, Palm Beach in 1951 (USNM 50381, U.S. National Museum of Natural History 2014), St. Lucie and Fort Pierce inlets of the Indian River Lagoon in the 1980s (Mook 1983), and Fernandina Beach in 2006 (FLMNH 7900, Florida Museum of Natural History 2014). Specimens reported from Georgia and South Carolina (1980-2006) were collected in 30-35 m of water on the Continental Shelf (DeVictor and Morton 2010; U.S. National Museum of Natural History 2014), where Gulf Stream waters may have moderated winter temperatures.

Invasion History on the Gulf Coast:

Carijoa riisei was first reported from the Gulf Coast in Dry Tortugas, Florida in 1869 (YPM IZ 002392 CN, Yale Peabody Museum 2014). It has been reported from Texas to the tip of Florida, often 30-100 km offshore, at depths of 13 to 45 m (U.S. National Museum of Natural History 2014). One collection was made from a jetty in 0–8 m of water in Port Isabel, Texas in 1960 (USNM 51966, U.S. National Museum of Natural History 2014). Another collection was made west of the Florida Keys at 732 m depth in 1962 (USNM 93253, U.S. National Museum of Natural History 2014). The overall abundance of C. riisei relative to native soft corals in the Gulf of Mexico is unclear.

Invasion History in Hawaii:

Carijoa riisei was first found in the Hawaiian islands in a cave off the leeward coast of Oahu in 1966, and then found in Pearl Harbor, in fouling communities in 1972 (Carlton and Eldredge 2009; Concepcion et al. 2010). By 2006, it was found throughout the Hawaiian Islands, especially in shaded areas with abundant plankton and moderate currents (Coles et al. 1999a; Coles et al. 2002a; Coles et al. 2004; Carlton and Eldredge 2009). Initially, C. riisei was regarded as an inhabitant of fouling communities in disturbed, shallow-water areas, but in 2001, it was found at depths up to 110 m, overgrowing beds of commercially valuable Black Coral (Antipathes dichotoma) (Grigg 2003; Grigg 2004; Kahng and Grigg 2005). Genetic studies indicate that this coral was introduced to Hawaii from the Indo-Pacific, separately from its introduction to the Caribbean (Concepcion et al. 2010).

Invasion History Elsewhere in the World:

Carijoa riisei has an early invasion history in the Caribbean and Western South Atlantic (Concepcion et al. 2010). It was first described from the Virgin Islands in 1860 (Deichmann 1936). In Brazil, it was collected in Rio de Janeiro Bay before 1867 (Müller 1867, as C. rupicola, DeVictor and Morton 2010) and Bahia in 1876 (USNM 50377, U.S. National Museum of Natural History 2014). It was collected in Kingston, Jamaica in 1885 (USNM 7541, U.S. National Museum of Natural History 2014). The coral is abundant in shallow coastal waters throughout the Caribbean, and south to Florianopolis, Brazil (Lira et al. 2009; Castro et al. 2010; Barbosa et al. 2014). It has also been collected in depths up to 309 and 508 m (Castro et al. 2010; DeVictor and Morton 2010). Carijoa riisei has also been collected off western Africa, on the island of Sao Tome (Concepcion et al. 2010), and on oil platforms off Gabon  (Friedlander et al. 2014).

Carijoa riisei appears to be a recent invader on reefs, on the Pacific coast of Colombia, where it was first noticed by divers around 2000 at Ensenada de Utría, Punta, and on Isla Malpelo. This coral seems to be rapidly overgrowing and replacing native octocorals (Sanchez et al. 2014). Genetic studies indicate that this population is closely related to introduced populations in the Caribbean, possibly transported through the Panama Canal (Quintanilla et al. 2017). It has been found  on fouling plates on Cocos Island, Costa Rica (Ruiz et al.  2021, unpublished).

Owing to its reported Caribbean origin, C. riisei has been reported as an invader in Indonesia (Calcinai et al. 2004) and India. Indian locations include the Andaman Islands (Divya et al. 2012), the Gulf of Mannar (Padmakumar et al. 2011), and the Gulf of Kutch (Yogesh Kumar et al. 2014). Local range extensions are possible within the Indo-Pacific range, but these records may result from collecting in poorly studied areas.

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Description

Carijoa riisei is a soft coral belonging to the subclass Octocorallia. As the name of the subclass suggests, the polyps have eight pinnate tentacles and eight complete mesenteries, dividing their gastrovascular cavities. The polyps grow in a colony connected by a mass of tissue called coenenchyme, which consists of mesoglea, perforated by gastrodermal tubes which connect the polyps' interiors. The mesoglea contains calcareous skeletal particles, called sclerites, which form an internal skeleton for the colony (Barnes 1983, DeVictor and Morton 2010). Colonies of C. riisei grow by monopodial branching from a single axial zooid, in which the branches arise from a single stem, which progressively lengthens. Colonies may be as much as 355 mm high. The colonies are connected by creeping stolons, which give rise to new colonies. The stem is smooth near its base, but in its upper ridges has eight longitudinal ridges. The colonies are bushy and densely branched. Zooids sometimes arise in pairs from a branch, or may be spiral, with four zooids in each turn (Deichmann 1936; Bayer 1981; Castro et al. 2010; DeVictor and Morton 2010; Barbosa et al. 2014). Polyps are cylindrical, and the external portion (anthocodium) can fully retract into the portion of polyp (anthostele) which is embedded in the coenenchyme. The sclerites are stick-like in form, often branching, with thorny projections, and often occur in tangled masses. The polyps are white, while the branches vary from white to orange or reddish brown (Deichmann 1936; Bayer 1981; Castro et al. 2010; DeVictor and Morton 2010).

Concepcion et al. (2008; 2010) treat most of the Carijoa specimens in their genetic analysis as C. riisei, although they have excluded a transparent morph found in Hawaii, as possibly a cryptic species. However, they acknowledge that the taxonomy of this coral is unresolved.

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Taxonomy

Ecology

General:

Carijoa riisei is known from a variety of tropical and subtropical marine habitats over a wide depth range. Its range of habitats include rocky offshore reefs, coral reefs, caves, crevices, mangrove roots, and from docks, piers, offshore oil platforms, and ship hulls (Carlton and Eldredge 2009; Lira et al. 2009; Friedlander et al. 2013; Barbosa et al. 2014; U.S. National Museum of Natural History 2014). It can occur from the low tide line to 700 m depth. It has been collected in warm-temperate habitats off Georgia and South Carolina, but at depths of 30-35 m (DeVictor and Morton 2010), where the Gulf Stream may moderate winter temperatures. In deep water off Maui, C. riisei appeared to be limited by an isotherm of 22°C (Kahng and Grigg 2005). Precise salinity tolerances are not known, but this coral survived only two days at 25 PSU, and was killed within minutes by fresh water (Toonen et al. 2007). This coral is often found in turbid waters with swift currents. The polyps feed by extending their tentacles in the current and trapping particles. In Brazil, 88% of the ingested biomass was phytoplankton, mostly cyanobacteria and diatoms, while 7% was zooplankton, and 5% was unidentified (Lira et al. 2009). Two specialized predators, the nudibranchs Phyllodesmium poindimiei and Tritoniopsis elegans, native to the Indo-West Pacific, appeared in the Hawaiian Islands in 2005 (Carlton and Eldredge 2009; Wagner et al. 2009). Phyllodesmium poindimiei appears to feed on this coral exclusively, while T. elegans is a generalist (Wagner et al. 2007; Carlton and Eldredge 2009; Wagner et al. 2009). Otherwise, C. riisei appears to have few predators in Hawaiian or Atlantic waters (Concepcion et al. 2010).

Food:

Phytoplankton, zooplankton

Consumers:

Phyllodesmium poindimiei (nudibranch)

Trophic Status:

Suspension Feeder

SusFed

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Habitats

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

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Life History

Carijoa riisei colonies, like those of other octocorals, produce new polyps though asexual reproduction. Unlike scleractinian corals, the polyps in a colony are gonochoristic, almost all of a single sex, either male or female. Hermaphroditic colonies comprised 1.3% of specimens examined in Hawaii. Hermaphrodites may represent colonies in the process of changing sex (Kahng et al. 2008). Colonies first start producing sperm or oocytes at 25–54 cm height (Kahng et al. 2008; Barbosa et al. 2014). Female polyps in Brazil produced a mean of 3.5 oocytes per polyp, while Hawaiian polyps produced 7.4. Sexual reproduction in both locations is continuous throughout the year. Fertilized eggs produce planula larvae, which are not brooded. The duration of the planular stage and the potential for dispersal is not known (Kahng et al. 2008; Concepcion et al. 2010). In Pacific Panama, numbers and size of eggs increased during the upwelling season (Quintero-Arrieta et al. 2023).

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	23	Field, based on depth distribution off Maui (Kahng and Grigg 2005)
Minimum Reproductive Temperature	None	Subtropical-Tropical
Broad Salinity Range	None	Euhaline

General Impacts

Carijoa riisei has long been regarded as a native species in the Western Atlantic, so its impacts on native corals or other reef and fouling community inhabitants have not been studied. In Hawaii, it was initially regarded as a relatively benign fouling organism, often in disturbed harbors and bays (Coles and Eldredge 2002; Carlton and Eldredge 2009). However, its invasion of a deep-water bed of Black Coral (Antipathes dichotoma and A. grandis) at 30–110 m depth, off Hawaii, resulted in extensive mortality of the native species. This invasion had both ecological and economic implications, since the Black Coral is commercially harvested for jewelry (Grigg et al. 2005; Kahng and Grigg 2005).  However, on shipwrecks off Pernambuco, Brazil, formed extensive three-dimensional structures which greatly increased the diversity  and abundance of epibenthic fauna., and so is considered an ecosystem engineer (Feitosa de Padua et al. 2023).  Carijoa riisei  is considered a serious competitor to native corals in Pacific Panama (Quintero-Arrieta et al. 2022).

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Regional Impacts

SP-XXI	None		Ecological Impact	Competition
Initially, Carijoa riisei was known as an increasingly abundant fouling organism in shallow waters, often on man-made structures in disturbed habitats, with little impact on native corals (Coles and Eldredge 2002). However, a survey of native Black Coral (Antipathes dichotoma and A. grandis) at 30-110 m depth found that many of the corals were being overgrown and killed by the invading Snowflake Coral. In a 2001 survey, 90% of the Black Corals had been overgrown and killed (Grigg 2003; Kahng and Grigg 2005). Exposure to fresh water was tested as an eradication method. Exposure killed polyps in 1.5 minutes, while 15 PSU water took 90 minutes, and 25 PSU water took 2 days. Use of low-salinity water would be limited mostly to man-made structures, possibly by wrapping pilings in plastic (Toonen et al. 2007).
SP-XXI	None		Economic Impact	Fisheries
The Black Coral fishery (Antipathes dichotoma and A. grandis) in the Hawaiian Islands was valued at about ~$30 million per year (Grigg et al. 2004). The invasion by Carijoa riisei on beds of Black Coral off Maui, combined with intensified harvesting, has contributed to a ~25% decrease in biomass of the corals (Grigg et al. 2004). These corals are commercially harvested for the manufacture of coral jewelry. Grigg et al. (2004) recommend changes in harvest quotas and size limits to adjust to increased mortality and decreased recruitment of the Black Corals. Limits were imposed on the Hawaiian Black Coral harvest in 2008 (Federal Register 8/13/2008).
SEP-I	None		Ecological Impact	Competition
In 2008-2012, rapid overgrowth of native octocorals on reefs in Pacific Colombia, by C. riisei, has been observed (Sanchez and Ballesteros 2014).
SEP-I	None		Ecological Impact	Habitat Change
None
SA-II	None		Ecological Impact	Competition
Areas dominated by Carijoa riisei (Snowflake Coral) resisted colonization by the Sun-Coral (Tubastraea coccinea , in at Búzios Island, Sao Paulo, Brazil (Mizrahi et al. 2017).
SA-III	None		Ecological Impact	Habitat Change
Colonies of Carijoa riisei on shipwrecks off Pernambuco, Brazil, formed complex three-dimensional structures, and supported an increased abundance (10X) and diversity (1.5X) of epibenthic invertebrates.
HI	Hawaii		Ecological Impact	Competition
Initially, Carijoa riisei was known as an increasingly abundant fouling organism in shallow waters, often on man-made structures in disturbed habitats, with little impact on native corals (Coles and Eldredge 2002). However, a survey of native Black Coral (Antipathes dichotoma and A. grandis) at 30-110 m depth found that many of the corals were being overgrown and killed by the invading Snowflake Coral. In a 2001 survey, 90% of the Black Corals had been overgrown and killed (Grigg 2003; Kahng and Grigg 2005). Exposure to fresh water was tested as an eradication method. Exposure killed polyps in 1.5 minutes, while 15 PSU water took 90 minutes, and 25 PSU water took 2 days. Use of low-salinity water would be limited mostly to man-made structures, possibly by wrapping pilings in plastic (Toonen et al. 2007).
HI	Hawaii		Economic Impact	Fisheries
The Black Coral fishery (Antipathes dichotoma and A. grandis) in the Hawaiian Islands was valued at about ~$30 million per year (Grigg et al. 2004). The invasion by Carijoa riisei on beds of Black Coral off Maui, combined with intensified harvesting, has contributed to a ~25% decrease in biomass of the corals (Grigg et al. 2004). These corals are commercially harvested for the manufacture of coral jewelry. Grigg et al. (2004) recommend changes in harvest quotas and size limits to adjust to increased mortality and decreased recruitment of the Black Corals. Limits were imposed on the Hawaiian Black Coral harvest in 2008 (Federal Register 8/13/2008).

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