Pterois volitans

Invasion History

First Non-native North American Tidal Record: 1985
First Non-native West Coast Tidal Record:
First Non-native East/Gulf Coast Tidal Record: 1985

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General Invasion History:

North American Invasion History:

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Description

Pterois volitans (Red Lionfish) has an almond-shaped body, and a large head, one-third to one-half body length, with numerous head spines, and often with large tentacles above the eyes. Dorsal fin spines and rays of dorsal, anal, pelvic and pectoral fins are greatly elongated, making the fish look a little like an exploding firework. The caudal fin is rounded. Eleven species of Lionfish (Pterois are known, some very similar morphologically to P. volitans. Pterois miles, very similar to P. volitans, has also been introduced to the the Western Atlantic, and can be distinguished only by meristic counts, morphometrics, and fine morphological details, or molecular methods (Schultz 1986; Hamner et al. 2007; Freshwater er al. 2009; Florida Museum of Natural History 2017; Froese and Pauly 2017; USGS Nonindigenous Aquatic Species Program 2017).

The dorsal fin of Pterois volitans has 13 spines and 9-12 (usually 11) soft rays. The anal fin has 3 spines, and 6-8 (usually 7) soft rays. (For P. miles, the usual numbers are 10 dorsal rays and 6 anal rays). Color patterns vary greatly with environment, but are basically zebra-striped in red and white. The soft dorsal, anal, and caudal fins are spotted. (Pectoral fins and fin spots are smaller in P. miles). The usual maximum size of P. volitans is ~380mm, but one specimen off Venezuela reached 457 mm (Schultz 1986; Ehemann 2017; Florida Museum of Natural History 2017; Froese and Pauly 2017; USGS Nonindigenous Aquatic Species Program 2017).

Of 180 specimens of Western Atlantic Pterois examined at the molecular level by Hamner et al. 2004), 168 were identified as P. volitans, primarily a western Pacific species, 12 specimens as P. miles, whose range encompasses thee wstern tropical/subtropical Indian Ocean (Africa-Red Sea-Australia) (Hamner et al. 2007). For both species, intraspecific variation in color and morphology is greater than that between species (Schultz 1986; Lowe 2016). A dramatic case is the appearance and rapid spread of a an apparently distinct morphoology of the supraoptical tentacle in both species, both in Indo-Pacific and Atlantic populations. The supraoptical tentacle can either be a simple, unbranched tentacle, or have a highly branched 'peacock-feather' morphology. This polymorphism probably evolved before the divergence of the two species (Morris and Freshwater 2008). Recent genetic studies indicate that hybridization has been occurring between P. volitans and P. miles in the Atlanitc, and P. volitans, in its native Indo-Pacific, has undergone genetic exchange with the more localized species P. lunulata (Luna Lionfish) and P. russelli (Plaintail Turkeyfish) (Wilcox et al. 2018). These authors suggest that heterosis ('hybrid vigor') may have contributed to the invasiveness of inttoduced lionfishes.

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Taxonomy

Ecology

General:

Red Lionfish (Pterois volitans) have separate sexes, and spawn in pairs, after a courtship ceremony. Male lionfish mature at about 100 mm, and females at about 180 mm. Lionfish are synchronous, ineterminant spawners. The ovaries contain eggs in all stages of development, and spawning can take place year-round. Mature eggs have a mean diameter of 804 um, Eggs are released in a mass, with an estimated 24,630 eggs, surrounded by a ball of mucus. Egg masses are released, two at a time, near the surface (Morris et al. 2009; Morris and Whitfield 2009; Morris et al. 2011; Gardner et al. 2015). In North Carolina, Lionfish spawn about every four days, resulting in an annual fecundity of about two million eggs per year (Morris and Whitfield 2009). The larvae remain in the plankton for 20-35 days, as estimated from daily otolith rings (Ahrenholtz and Morris 2010). The planktonic larvae hatch with a yolk-sac, which is resorbed, and then feed on plankton. The offshore spawning locations, larval period, and pelagic mode of development permit long-distance dispersal by currents (Morris and Whitfield 2009; Morris et al. 2009; Vasquez-Yeomans et al. 2011; Kitchens et al. 2017). Settlement probably takes place predominantly but not exclusively in shallow habitats, including mangroves, seagrasses, estuaries, and around man-made structures, where juvenile lionfish are more common. Larger fish tend to predominate in deep-reef habitats (Biggs and Olden 2011; Claydon et al. 2012; Jud et al. 2011; Cure et al. 2014).

The Red Lionfish is generally regarded as a strictly marine, and tropical fish, but it is highly mobile and versatile, capable of using a wide range of habitats. In experiments where temperature was decreased gradually, the mean lethal temperature was 10 C, while feeding ceased at 16 C. In the field, populations off NC, were limited by a 12 C isotherm (Kimball et al. 2004), permitting the fish to winter in Gulf Stream-influenced waters (Kimball et al. 2004). The mean temperature for Lionfish in Onslow Bay, North Carolina was 15.3 C, at a depth of 38-46 M (Whitifield et al. 2014). Upper lethal temperature, determined as a critical thermal maximum (CTM), was 34-39 C (Aiken et al. 2014). Juvenile and adult lionfish were found through much of the Loxahatchee estuary, Florida (Jud et al. 2011). Lionfish survived 28 days at 7 PSU with no mortality or changes in behavior, feeding and growth. Fish in the Loxahatchee estuary occasionally occurred at salinities as low as 1 PSU, but did not survive prolonged exposure to these conditions (Jud et al. 2011; Schofield et al. 2015). Lionfish utilize a range of habitats, including coral reefs, mangroves, seagrass beds, and man-made structures, such as pilings (Barbour et al. 2010; Biggs and Olden 2011; Claydon et al. 2012; Jud et al. 2011; Pimiento et al. 2013). Lionfish have been widely reported in shallow water, but also from the mesophotc zones at the base of reefs, and from deeper slope waters at 200-300 m depth (Gress et al. 2017). The visual system of lionfishes are similar to those of Western Atlantic estuarine piscivores, but they are more sensitive to hypoxia, which could limit their penetration into estuaries (Hasenei et al. 2020).

Juvenile and adult Lionfishes are carnivorous. In the Loxahatchee River estuary, where juvenile fish predominated, the prey were about half-and-half fishes and crustaceans (crabs and shrimps). By weight, penaeid shrimps, blennies, and Gerreidiae (mojarras, a small perch-like fish) predominated (Jud and Layman 2011). On reefs in the Bahamas, Côté and Maljkovic (2010) observed Lionfish consuming many species of reef fishes. Lionfishes hunt throughout the diel cycle, but feeding rates are highest near dawn and sunset, and on overcast days (Côté and Maljkovic 2010; Green et al. 2011). In experiments, feeding rates were higher under white or blue light than red light, and lowest in darkness. Attack rates were much higher in environments of low habitat complexity (South et al. 2017).

Lionfishes' long-venomous spines limit predation, but they are eaten by native Nassau Groupers (Epinephelus striatus) (Maljkovic and Van Leeuwen 2008; Mumby et al. 2011). Lionfish can be eaten by humans, when the spines are removed, and are highly regarded as food in Asia. Fisheries are being encouraged in the Caribbean, as a possible means of population control (Morris et al. 2012). Introduced populations of lionfish have a much lower incidence of parasites than populations in the naitve range (Tuttle et al. 2017). Lionfish are not social or schooling, but do aggregate in structurally complex habitats. This might aid the development of artificial structures for the attraction and acpture of Lionfish (Hunt et al. 2019).

Food:

Crustaceans (shrimps), Fishes, Crabs

Consumers:

Humans; fishes

Trophic Status:

Carnivore

Carn

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Habitats

General Habitat	Coral reef	None
General Habitat	Unstructured Bottom	None
General Habitat	Rocky	None
General Habitat	Marinas & Docks	None
General Habitat	Coarse Woody Debris	None
Tidal Range	Subtidal	None
Vertical Habitat	Epibenthic	None
Vertical Habitat	Nektonic	None

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Life History

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	9.5	Minimum CT min (Barker et al. 2017)
Maximum Temperature (ºC)	39.5	Aikins et al. 2014 Critical Thermal Maximum, Aikens et al. 2014; Barker et al. 2017 http://www.fishconserve.org/arcel/admin/uploads/FPLP112.pdf
Minimum Salinity (‰)	7	Experimental (Jud et al. 2014; Schofield et al. 2015). Long-term survival and growth occurred at 10-34 PSU (Schofield et al. 2015), but lionfish tolerated short exposures (>7 days) at 4-5 PSU (Jud et al. 2014; Schofield et al. 2015).
Maximum Salinity (‰)	36	Typical Gulf of Mexico salinity
Minimum Duration	20	Approximate time from hatching to settlement, based on daily otolith rings, Bahamas (Ahrenholz and Morris 2010)
Maximum Duration	35	Approximate time from hatching to settlement, ased on daily otolith rings, Bahamas (Ahrenholz and Morris 2010)
Minimum Length (mm)	100	Minimum for males, 170 mm for females (Morris and Whitfield 2019); Morris et al. 2011
Maximum Length (mm)	467	Bermuda (Eddy et al. 2019); 457 mm Ehemann (2017 , One specimen, Venezuela)
Broad Temperature Range	None	Subtropical-Tropical
Broad Salinity Range	None	Mesohaline-Euhaline

General Impacts

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Regional Impacts

CAR-V	None		Ecological Impact	Predation
In transplantation experiments, patch reefs with a transplanted lionfish had reduced recruitment of juvenile native fishes. Juvenile native fishes were found in the lionfishes' gut contents (Albins and Hixon 2008; Albins et al. 2017). Mortality rates in a small fish, Gramma loreto (Fairy Basslet) greatly increased after the lionfish invasion, compared with a pre-invasion experiment (Ingeman and Webster 2015). In later experiments, in which prey and lionfish densities were maipulated, some low-density basslet populations were extirpated by lionfish predation (Ingeman et al. 2016). Isotopic values (15N, 13C) values varied, but shopwed an increasing trophic niche and diversity for large lionfish (Malpici-Cruz et al. 2019). Juveniles of the native Striped Parrotfish (Scarus iseri) have an unlearned response to Nassau Grouper (Epinephelus striatus but have to learn avoidance of Pterois spp. (Berchtold and Cote 2020).
CAR-V	None		Ecological Impact	Habitat Change
Predation by lionfish has decreased local populations of herbivorous fishes, resulting in increased macroalgal populations at deeper depths, resulting in declines of corals and sponges (Lesser and Slattery 2011).
CAR-V	None		Ecological Impact	Trophic Cascade
Predation by lionfish has decreased local populations of herbivorous fishes, resulting in increased macroalgal populations at deeper depths, resulting in declines of corals and sponges (Lesser and Slattery 2011). Predation by Lionfish resulted in reduced grazing by herbivorous fishes on algae, both through direct predation, but also by inducing behavioral changes, with reduced abundances of small fishes in the presence of lionfish (Kindinger and Albins 2016). Differential predation affected competitive relations between planktivorous fairy and blackcap basslets (Gramma loreto and Gramma melacara) (Kindinger 2018). Predation by lionfish affected the abundance of one abundant cleaner fish (Thalassoma bifasciatum, Bluehead Wrass), but did not affect another Elacatinus genie (Cleaner Goby). Cleaner fishes affect the health and social behavior of coral reef fish communities by removing parasites (Tuttle 2017),
CAR-IV	None		Economic Impact	Health
Ciguatera toxin was found in Lionfish in St. Maarten in 2011. and people were advised not to eat the fish (Associated Press 2011). Howver, test results on uncooked lionfish can be misleading, because they contain another group of toxins (Scorpaenitoxins), which give positive results, but are destroyed by cooking (Hixon et al. 2015). In Martinique, 117 patients in 2011-2014, stung by lionfish spines, were examined. All had local swelling. Other common symptoms were more extensive swelling, gastointestinal disorders, abdominal cramps, fainting, tachycardia, and hyptertension. Immersion of the stung area in warm water reduced pain duration and the chance of infection (Reisiere et al. 2015).
CAR-VII	Cape Hatteras to Mid-East Florida		Ecological Impact	Predation
The population increase in Pterois spp. (mostly P. volitans) from 1990 to 2014 has been associated with a 45% decline of a common native fish, Tomtate (Haemulon aurolineatum) (Ballew et al. 2016).
CAR-IV	None		Ecological Impact	Predation
Lionfish were found to be feeding on a newly described goby (Palatogobius incendius) living in deep-water mesophotic reef areas (Tornabene and Baldwin 2017).
CAR-III	None		Ecological Impact	Predation
Lionfish were found to be feeding on a newly described goby (Palatogobius incendius) living in deep-water mesophotic reef areas (Tornabene and Baldwin 2017)
S200	Biscayne Bay		Ecological Impact	Competition
A native predator, Cephalopholis cruentata consumed less prey in areas of high lionfish density, but condition of the native fish was not affected (Curtis et al. 2017).
CAR-I	Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida		Ecological Impact	Competition
A native predator, Cephalopholis cruentata consumed less prey in areas of high lionfish density, but condition of the native fish was not affected (Curtis et al. 2017).
CAR-I	Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida		Ecological Impact	Predation
Lionfishes in natural habitats and artificial reefs consumed a wide range of fish and invertebrate prey, with increasing piscivory in larger fish. Thirty-three species of prey were identified (Dahl and Patterson 304; Dahl et al. 2017). Modeling indicates that populations of several important grouper and snapper species (particularly Epinephelus morio Red Grouper) could be affected by high predicted abundances of lionfish off the Florida peninsula and Texas Gulf Coast (Johnston et al. 2017). Caging experiments in the Loxahatchee estuary of the Indian River Lagoon found that Lionfish caused a 90% reduction in Grass Shrimp (Palaemon) abundance, and a general reduction of benthic invertebrate abundance and diversity (Layman et al. 2014). Experimental studies and modeling indicate that P. volitians had a higher predation impact than two native predators (Epinephalus morio, Red Grouper; Cephalopholis cruentata, Graysby Grouper), because of the combination of high predator efficiency and high abundance (DeRoy et al. 2020).
CAR-I	Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida		Economic Impact	Fisheries
Modeling indicates that populations of several commercially important grouper and snapper species (particularly Epinephelus morio Red Grouper) could be affected by high predicted abundances of lionfish off the Florida peninsula and Texas Gulf Coast. Control methods should be focused on these areas (Johnston et al. 2017).
S190	Indian River		Ecological Impact	Predation
Caging experiments in the Loxahatchee estuary of the Indian River Lagoon found that Lionfish caused a 90% reduction in Grass Shrimp (Palaemon) abundance, and a general reduction of benthic invertebrate abundance and diversity (Layman et al. 2014).
NA-ET4	Bermuda		Ecological Impact	Predation
Lionfish in Bermuda preyed on a wide range of fishes and crustaceans (from 46 families), including ecologically important and economically improtant species (Eddy et al. 2017).
FL	Florida		Ecological Impact	Competition
A native predator, Cephalopholis cruentata consumed less prey in areas of high lionfish density, but condition of the native fish was not affected (Curtis et al. 2017).
FL	Florida		Ecological Impact	Predation
Caging experiments in the Loxahatchee estuary of the Indian River Lagoon found that Lionfish caused a 90% reduction in Grass Shrimp (Palaemon) abundance, and a general reduction of benthic invertebrate abundance and diversity (Layman et al. 2014).

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