Carijoa riisei

Invasion History

First Non-native Panama (Pacific) Tidal Record: 2013
First Non-native Panama (Caribbean) Tidal Record: 1935

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Panama Invasion History:

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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 It has been found  on fouling plates on Cocos Island, Costa Rica (Ruiz et al.  2021, unpublished).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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