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    Taxonomic name: Pterois volitans (Linnaeus 1758)
    Synonyms: Brachirus zebra (Quoy and Gaimard, 1825), Gasterosteus volitans, Pterois cristatus, Pterois geniserra, Pterois muricata, Pterois zebra Quoy and Gaimard, 1825, Scorpaena volitans (Linnaeus, 1758)
    Common names: butterfly cod (English-Christmas Island), cá Mao Tiên (Vietnamese-Viet Nam), chale (Swahili-Kenya), firefish (English), hana-minokasago (Japanese-Japan), Indo-Pacific red lionfish, laffe volant (French-Seychelles), lepu-penganten (Malay-Indonesia), lionfish (English), lionfish scorpion (English), ominokasago (Japanese-Japan), ornate butterfly-cod (English), ornate butterfly-cod (English-USA), peacock lionfish, poisson scorpion (French-Mauritius), poisson volant (French-Seychelles), poisson-dindon (French-Mauritius), red firefish (English-Australia and USA), red lionfish (English-New Zealand), sausau-lele (Samoan-Samoa), scorpion fish (English), scorpion volitans, skrzydlica pstra (Polish-Poland), turkey fish (English-Guam, Micronesia and USA), turkeyfish (English), volitan lion (English), zebrafish (English)
    Organism type: fish
    The Indo-Pacific red lionfish (Pterois volitans) is a beautiful but dangerous tropical fish that has spread to new marine environments through the aquarium trade. The lionfish has invaded the Northwestern Atlantic and the Caribbean in one of the most rapid marine finfish invasions in history. In some areas, it has the potential to displace commercially important species such as the grouper and reduce recruitment of juvenile fishes, which in turn disrupts marine ecosystem processes.
    Description
    Pterois volitans has elongated venomous dorsal fin spines and anal fin spines. It has 13 dorsal spines, 10 to 11 dorsal soft rays, 3 anal spines, 6 to 7 anal soft rays. Coastal populations are generally darker, sometimes almost black in estuaries (FishBase 2006). The membranes of fins are often spotted. The body is white or cream coloured with red to reddish-brown vertical stripes. The vertical stripes alternate from wide to very thin and sometimes merge along the flank to form a V (Schofield and Fuller 2006). The maximum length of an adult is 38cm (FishBase 2006) and the maximum body weight is 1.1kg to 1.2kg (Fishelson 1997). Reports of a 43cm individual have been obtained in its introduced range.
    Similar Species
    Pterois miles

    More
    Occurs in:
    marine habitats
    Habitat description
    Pterois volitans is a tropical marine fish which usually occurs in waters with temperatures of between 22°C and 28°C. Lower temperature ranges have been observed in the USA (14°C to 24°C) (Meister et al. 2005). The depth range of this species is 10 to 175 meters (Schofield & Fuller 2006). The red lionfish favours coral reefs and rocky outcrops (DaCosta-Cottam et al. 2009), although it has been observed over coral patches, near sandy bottoms and in mangrove, seagrass and even canal habitats (Schofield 2009; González et al. 2009). It is also found in lagoons and harbours (FishBase 2006). It hides in unexposed places during the daytime often with its head down, practically immobile.

    Richter (2009) hypothesises that the range of the lionfish will reach the Gulf of Mexico and expand south along the eastern coast of South America until it reaches a southern latitude where it cannot tolerate low temperatures (Hare and Whitfield 2003, in Richter 2009). The southernmost sighting has been in Columbia, though there is potential for the lionfish to extend past that (Schofield 2009, in Richter 2009). The northern limit of the range is North Carolina; a functional expansion northward is unlikely unless the lionfish population undergoes a rapid evolution to thermal tolerance; some juveniles have been seen as far north as Long Island Sound, although they cannot become established due to cold winters (Richter 2009). Temperature is a possible factor limiting; the mean chronic lethal minimum is 10°C and the mean temperature of feeding cessation is 16°C, indicating that the lionfish might over-winter on the southeast United States continental shelf, with a northern limit of Cape Hatteras (Kimball et al. 2004).

    General impacts
    For a detailed account of the environmental impacts of Pterois volitans please read: Pterois volitans (Indo-Pacific Red Lionfish) Impacts Information. The information in this document is summarised below. Invasive lionfish are a concern to coastal managers due to their potential threat to fisheries resources, native fish communities and human health (Morris et al. 2008).

    Ecosystem change: While few ecological studies have been conducted (but see Albins & Hixon 2008) it is clear that the lionfish’s presence in the Caribbean is a worrying one. Lionfish are highly piscivorous and reduce the recruitment of juvenile fishes, which in turn disrupts marine ecosystem processes and reduces reef biodiversity (Albins and Hixon 2008; Morris et al. 2008).

    Reduction in native biodiversity: If their populations are allowed to continue growing unchecked, lionfish have the potential to severely reduce reef biodiversity, with the possible extinction of several species; although it is still too early to be definitive, anecdotal evidence from the Bahamas corroborates this premise (Dell 2009).

    Predation: Albins and Hixon (2008) showed that lionfish can drastically reduce recruitment of native fishes on small patch reefs in the Bahamas. They are potentially capable of decimating indigenous reef fish populations in the Caribbean due to their lack of natural predators and voracious appetite (Valdez Mascari & Aguiar 2009).

    Competition: Not only do lionfish consume large quantities of juvenile fish (such as grouper and yellow-tail snapper) but they also out-compete native species (such as scamp, gag, and yellow-mouth grouper) for food (Morris et al. 2008; Dell 2009).

    Economic/Livelihoods: In addition, by reducing populations of commercially important species such as grouper (Albins and Hixon 2008) they may as a consequence damage the economy of island communities which are dependent on such fishing industries.

    Human health: Lionfish are venomous with their spines containing apocrine-type venom glands (Morris et al. 2008). Lionfish venom has been found to cause cardiovascular, neuromuscular, and cytolytic effects ranging from mild reactions such as swelling to extreme pain and paralysis in upper and lower extremities (Kizer et al. 1985, in Morris et al. 2008). The toxin in lionfish venom contains acetylcholine and a neurotoxin that affects neuromuscular transmission (Cohen and Olek 1989, in Morris et al. 2008). Lionfish spines can prove dangerous to divers, snorkelers and aquarium enthusiasts (Morris et al. 2008; Schofield 2009). Stings are not fatal, but intensely painful and often requiring hospitalisation (Morris et al. 2008). Lionfish stings can be treated by heating the afflicted part in hot water (to 45° C) for 30 to 90 minutes and applying corticoids to the area (FishBase 2006); medical attention should be sought immediately (Cayman Islands Government Undated).

    Uses
    Pet/aquarium trade: The red lionfish is a popular species among tropical fish enthusiasts (Schofield and Fuller 2006).
    Fisheries: Commercial fisheries of red lionfish exist in the lionfish’s native range (FishBase 2006) and it is reportedly a viable commercial species in its introduced range also, although encouraging such fisheries may encourage the intentional dispersal of the fish (see Management Information).
    Bioprospecting: Work by Sri Balasubashini and colleagues (2006a, 2006b, in Morris et al. 2008) indicated that lionfish (P. volitans) venom produces antitumor, hepatoprotective, and antimetastatic effects in mice suggesting a promising application for cancer research.
    Notes
    Lionfish comprises a species complex whose native range encompasses the Red Sea, the Indian Ocean and the western Pacific (Schultz 1986, in González et al. 2009). While traditional taxonomy indicates that P. volitans (Linnaeus 1758) and Pterois miles (Bennett 1828, in González et al. 2009) (devil firefish) comprise two different valid species (Schultz 1986, in González et al. 2009), recent molecular studies suggest that there is uncertainty in determining whether the two entities represent species or populations (Kochzius et al. 2003, Whitfield et al. 2007, in González et al. 2009). This species profile focuses on information on Pterois volitans; information on P. miles is included where appropriate.
    Geographical range
    Native range: Pterois volitans is widely distributed throughout the western Pacific and most of Oceania east to French Polynesia (Schofield and Fuller 2006).
    Known introduced range: P. volitans has invaded the Atlantic coasts of the USA and the Caribbean (Schofield and Fuller 2006; González et al. 2009). The areas that are currently most affected are the southeastern coast of the United States, Bermuda, and the Bahamas; this is due to lionfish establishment since before 2005 (Richter 2009). Pelagic juveniles expatriate over great distances explaining the lionfish's broad geographical range (FishBase 2006). Molecular analyses have shown that both lionfish taxa are present in the Western Atlantic, with P. volitans being more than one order of magnitude more common than P. miles (Hamner et al. 2007, in González et al. 2009). To view the progress of lionfish distribution in the Atlantic Ocean since 1992, please see this Animated Distribution Map created by the United States Geological Survey.
    Introduction pathways to new locations
    Natural dispersal: Dispersal of lionfish presumably occurs during the pelagic larval phase during which larvae can be dispersed across great distances; for example, lionfish eggs released in the Bahamas are capable of dispersing to New England via the Gulf Stream (Morris et al. 2008). Larval connectivity models for reef fishes provide insight into lionfish larvae dispersal and are valuable for predicting the spread of lionfish. For example, further lionfish dispersal into the lower Caribbean and the Gulf of Mexico seems imminent (Cowen et al. 2006, in Morris et al. 2008). Based on the rapidity of lionfish establishment along the U.S. East Coast and the Bahamas, lionfish establishment along the southern edges of Central America (Nicaragua, Costa Rica, and Panama), the Yucatan peninsula, and the western Gulf of Mexico is likely within a few years or less (Morris et al. 2008).
    Pet/aquarium trade: P. volitans is a popular aquarium fish; aquarium releases are the most probable source of introduction of this invasive fish to new locations (Whitfield 2002).
    Ship ballast water: Eggs and larvae of the red lionfish may be transported via ballast water (Whitfield 2002).


    Local dispersal methods
    Natural dispersal (local):
    Other (local): Lionfish are thought to have been introduced accidentally into some parts of the Atlantic Ocean following the destruction of an aquarium by Hurricane Andrew (IUCN 2009; Cayman Islands Government Undated).
    Management information
    For a detailed account of managment of Pterois volitans please read: Pterois volitans (Indo-Pacific Red Lionfish) Management Information. The information in this document is summarised below.

    Preventative measures: Where the lionfish has not yet become properly established, preventative controls may include education of fishermen and other locals and encouraging the public to kill the fish on-sight and record positional data (Richter 2009). Some countries have urged recreational divers and snorkelers to record locations of lionfish sightings; other governments and programs have given monetary incentives to fishermen to catch and record positional data of lionfish (REEF 2009, in Richter 2009).

    Monitoring: Determining the extent of the lionfish invasion is necessary for effective management (Richter 2009). Organizations such as the Reef Environmental Educational Foundation (REEF) and the United States Geographical Survey-Nonindigenous Aquatic Species (USGS-NAS) have collected data of GPS locations, dates, water depth, and locality description of lionfish sightings (Richter 2009).

    Biocontrol: Groupers, a known natural predator of the lionfish, could hold the key to controlling this invasive species, however, this remains uncertain and unfortunately Atlantic populations of grouper are suffering from over-fishing (IUCN 2009). There is hope that through the establishment of Marine Protected Areas (MPAs) sufficient levels of groupers will return, providing an effective and natural means to control lionfish (IUCN 2009).

    Physical: In order to control and manage the lionfish invasion, culling programs have been introduced in the Cayman Islands, Bermuda and the Bahamas (4th UK CBD Report 2009; Adam-Whitmore 2009).

    Integrated Pest Management: Although bio-economic evaluations are necessary to determine the most effective management scheme, Richter (2009) suggests the establishment of a lionfish fishery may prove to be effective (Richter 2009). With the advent of an invasive species fishery, there are certain precautions, however; with the introduction of a potentially economically viable fishery, there lies the risk that people who reap the economic benefits may seek to maintain wild populations of the lionfish (Carlton, Pers. Comm., in Richter 2009).

    Nutrition
    Pterois volitans is an ambush predator that preys on small and juvenile fish, crustaceans (crabs and shrimps), molluscs and isopods. Lionfish are relatively quick to adapt to novel prey types, and quickly learn to avoid toxic prey (Fishelson 1997); in its invaded system in Caribbean reef ecosystems it uses its oversized, ornate pectoral fins to herd and ambush small reef fishes and crustaceans (Albins & Hixon 2008; Albins 2008). Analysis of gut contents and aquarium feeding trials confirm that the lionfish preys on a wide variety of native animals, including bony fishes and crustaceans, and that lionfish are capable of consuming large numbers of prey and large-sized prey relative to their body size (Albins 2008). For example, analysis of gut contents of 291 lionfish caught off the coast of Quintana Roo, Mexico, revealed the following prey items: bluehead wrasse, yellowhead wrasse, clown wrasse, creole wrasse, lizard fish, filefish, cardinal fish, blue chromis, brown chromis, fairy basslet, peppermint basslet, masked goby, goldspot goby, glass goby, rusty goby, spotted goatfish, trumpet fish, Spanish hogfish, spotted goatfish, slender filefish, redband parrot fish, hamlet fish, octopus, banded coral shrimp and night shrimp (D. Ponce-Taylor Pers. Comm.; Y.G. Hernandez Pers. Comm.). Assessments of invasive lionfish feeding suggest that lionfish are largely piscivorous, but also feed on a number of crustaceans; the particular taxa of highest importance in invasive lionfish diet will likely vary by habitat type and prey availability (Morris et al. 2008). A diet study of Bahamian lionfish was conducted by Morris and Akins (In Press) that showed that lionfish primarily eat teleost fishes (they documented 41 species in 21 families) as well as some crustaceans (about 15% of the diet by volume) (in Schofield 2009).

    In the Red Sea, individuals of the closely related P. miles were reported to feed on assorted taxa of benthic fishes including damselfish, cardinal fish, and anthias (Fishelson 1975 1997, in Morris et al. 2008). However, in the Pacific Ocean, P. lunulata were observed to feed primarily on invertebrates including penaeid and mysid shrimps (Matsumiya et al. 1980, Williams and Williams 1986, in Morris et al. 2008).

    Lionfish stomachs can expand over 30 times in volume when consuming a large meal; lionfish are capable of long-term fasting and are able to withstand starvation for periods of over 12 weeks without mortality (Fishelson 1997, in Morris et al. 2008). Lionfish (ranging from 30 to 300g) consume approximately 2.5% to 6% of their body weight per day (at 25°C to 26°C) in their native range; adults (ranging from 300 to 400 g) consume approximately 8.5 g of prey per day, which translates to 230 kg per year for 80 adult fish on a 1 kilometer stretch of coral reef (Fishelson 1997). Preliminary observations suggest that lionfish in their invaded range consume piscine prey at rates greater than in their native range (Morris et al. 2008).

    Reproduction
    Lionfish reach sexual maturity at around 18cm or 140g to 160g (approximately 1 to 2 years of age); they are external fertilizers that produce a pelagic egg mass following a courtship and mating process (Fishelson 1997; Florida Museum of Natural History 2006; DaCosta-Cottam et al. 2009). Reportedly females may produce up to 30 000 eggs a month, each of which will hatch in four days. The Pteroines, including P. miles and P. volitans, are gonochoristic; males and females exhibit minor sexual dimorphism only during reproduction (see Fishelson 1975, in Morris et al. 2008). Lionfish courtship has been well described by Fishelson (1975, in Morris et al. 2008) who provided a detailed description for the pigmy lionfish (Dendrochirus brachypterus) and reported similar courtship behaviors for Pterois spp. According to Fishelson, lionfish courtship, which includes circling, side winding, following, and leading, begins shortly before dark and extends well into nighttime hours; following the courtship phase, the female releases two buoyant egg masses that are fertilized by the male and ascend to the surface; the eggs and later embryos are bound in adhesive mucus that disintegrates within a few days, after which the embryos and/or larvae become free floating.

    P. miles and P. volitans ovarian morphology is similar to that reported for D. brachypterus (Fishelson 1978, in Morris et al. 2008) in that these fishes exhibit cystovarian type ovaries (Hoar 1957, in Morris et al. 2008) with ooctyes developing on stalks or peduncles. The oocytes are terminally positioned near the ovary wall, which secretes the encompassing mucus shortly before spawning.

    The seasonality of lionfish reproduction throughout their native range is unknown; invasive lionfish collected off North Carolina and in the Bahamas suggests that lionfish are reproducing during all seasons of the year (Morris et al. 2008).

    Lifecycle stages
    Larval stage descriptions for Pterois miles and P. volitans are incomplete with only one report by Imamura and Yabe (1996, in Morris et al. 2008) describing five P. volitans larvae collected off northwestern Australia. Scorpaenid larvae exhibit two morphologically distinct groups characterized as “morph A” and “morph B”; pteroine larvae are grouped among the morph B morphotypes, whose traits include: large head, relatively long and triangular snout, long and serrated head spines, robust pelvic spine, and pigment confined to the pectoral fins and postanal ventral and dorsal midlines (Leis and Rennis 2000, Washington et al. 1984, in Morris et al. 2008). Pterois sp. meristic characters are reported as 12 to 13 dorsal spines, 9 to 12 dorsal rays, three anal spines, 5 to 8 anal rays, 12 to 18 pectoral rays, one pelvic spine, five pelvic rays and 24 vertebrae (Imamura and Yabe 1996, Leis and Rennis 2000, in Morris et al. 2008).

    The size of P. miles or P. volitans larvae at hatching is unmeasured, but is likely to be approximately 1.5 mm based on reports for P. lunulata (Mito and Uchida 1958, Mito 1963, in Morris et al. 2008). The specific planktonic larval duration of lionfish is also unknown, although it is estimated to be between 25 to 40 days based estimates for Scorpaena (Laidig and Sakuma 1998, Hare and Whitfield 2003, in Morris et al. 2008).

    Compiled by: National Biological Information Infrastructure (NBII) & IUCN/SSC Invasive Species Specialist Group (ISSG)
    Updates with support from the Overseas Territories Environmental Programme (OTEP) project XOT603, a joint project with the Cayman Islands Government - Department of Environment
    Last Modified: Tuesday, 10 August 2010


ISSG Landcare Research NBII IUCN University of Auckland