| Common names: grouper (English), cabrilla (Espanol), mero (Espanol) |
Epinephelus itajara (Lichtenstein, 1822)
Goliath grouper, Itajara grouper
Body robust and oblong; wide and flat between eyes; eyes small; edge of preopercle angular; finely serrated; 21-24 gill rakers; dorsal fin XI, 15-16, the spines shorter than the rays; anal rays III, 8; pectoral rays 19; tail rounded.
Large fish: grey or greenish with pale blotches and smaller dark brown or blackish spots scattered over upper parts of head, body, and on pectoral fins. Small fish (< 100 cm) greenish to tawny brown with oblique, irregular darker brown bars.
The largest bony reef fish in the region, growing to 250 cm and 320 kg.
Size: 250 cm. Reaches at least 309 kg.
Habitat: rocky reefs as well as bays, estuaries and mangrove areas.
Depth: 1-100 m.
Western and eastern Atlantic and eastern Pacific; the SW and central E Gulf of California to Peru, the Revillagigedos, Cocos and Malpelo.
Global Endemism: All species, TEP non-endemic, East Pacific + Atlantic (East +/or West), Transisthmian (East Pacific + Atlantic of Central America), East Pacific + all Atlantic (East+West)
Regional Endemism: All species, Eastern Pacific non-endemic, Tropical Eastern Pacific (TEP) non-endemic, Continent + Island (s), Continent, Island (s)
Climate Zone: North Temperate (Californian Province &/or Northern Gulf of California), Northern Subtropical (Cortez Province + Sinaloan Gap), Northern Tropical (Mexican Province to Nicaragua + Revillagigedos), Equatorial (Costa Rica to Ecuador + Galapagos, Clipperton, Cocos, Malpelo), South Temperate (Peruvian Province )
Regularity: Regularly occurring
Type of Residency: Year-round
Global Range: Tropical and subtropical waters of the Atlantic and eastern Pacific oceans. Ranges from Florida to southern Brazil in the western Atlantic and is caught widely in the Gulf of Mexico and most of the Caribbean. Occurs from the Gulf of California to Peru in the eastern Pacific (Beebe and Tee-van 1933, Heemstra and Randall 1993, Greenfield and Thomerson 1997, Hoese and Moore 1998). Also occurs in east Africa from Senegal to the Congo, but reported as EPINEPHELUS ESONUE (Heemstra and Randall 1993). Rare in Aruba, Bonaire, and Curacao, and cited as uncommon in the Lesser Antilles such as Saba, St. Martin, and St. Eustatius (Nagelkerken 1981a, FAO 1993). Specific location records include: Bermuda (Smith 1958), Bahamas (Smith 1971, Roe 1976), Campeche Bank, Mexico (Roe 1976), eastern Gulf of Mexico or west Florida shelf (Tabb and Manning 1961, Smith et al. 1975, Colin 1990), Florida Keys (Starck, 1968; Voss et al. 1969), Cuba (Smith 1971), Jamaica (Thompson and Munro 1983), Hispaniola (Smith 1971), Brazil, and Colombia (Smith 1971). The range in the Atlantic is principally confined to south Florida (Huntsman et al. 1990); spawning aggregations reported only in the eastern Gulf of Mexico (Bullock and Smith 1991, Bullock et al. 1992).
Smith (1971): head broad and flat, large species, XI dorsal spines, vertical fins rounded, dorsal spines short, and small black spots.
Inshore/Offshore: Inshore, Inshore Only
Water Column Position: Bottom, Bottom only
Habitat: Reef (rock &/or coral), Rocks, Reef and soft bottom, Reef associated (reef + edges-water column & soft bottom), Soft bottom (mud, sand,gravel, beach, estuary & mangrove), Sand & gravel, Estuary
FishBase Habitat: Reef Associated
Habitat and Ecology
Found from inshore to about 100 m in reef, mangrove, seagrass, and estuarine habitats (Sadovy and Eklund 1999).
Juveniles live in shallow bays, holes, below undercut ledges in swift tidal creeks draining mangrove swamp, rivers and estuaries while adults live around structures in, near, and offshore (Bullock et al. 1992, Gerber et al. 2005, Koenig et al. 2007). Juveniles exhibit high site fidelity to mangrove habitat for 5-6 years, then emigrate to offshore reefs at body length of about 1 m TL (Koenig et al. 2007).
Juvenile distribution in mangroves depends on local water quality, particularly dissolved oxygen content (>4 ppm) and mid-range salinities (>10 ppt) (www.bio.fsu.edu/coleman_lab/goliath_grouper.html, accessed on 31st Dec 2005).
During a survey of the freshwater fish of southern Florida from 1976 to 1983, no E. itajara was collected although the salinity-tolerant juveniles could be found in shallow, costal waters (Loftus and Kushlan 1987). In 181 sites, presence of mangrove areas appears to be important for juveniles (Sadovy and Eklund 1999). Koenig et al. (2007) demonstrated the high nursery value of mangrove to juveniles.
The species feeds on a wide diversity of fishes and invertebrates (Sadovy and Eklund 1999). It is a classic apex predator, large, rare and only a few individuals occur on any given reef unit (Huntsman et al. 1999)
Up to 100, sometimes more, individuals aggregate to spawn at specific times and locations. The aggregations last only a few weeks each year and represent most of the total annual reproductive effort (Sadovy and Eklund 1999). Its reproductive season occurs between June and December, with peak activity indicated from July through September in the eastern Gulf of Mexico (Bullock et al. 1992). The species is one of the few groupers that aggregate in relatively shallow (10-50 m) water (Coleman et al. 2002).
Possible courtship activities (without spawning) were observed on a wreck in the eastern Gulf of Mexico at 33 m depth in August 1990 (Colin 1994). Wrecks are often noted to be spawning areas for this species.
A study indicated that a 1,322 mm standard length (SL) and a 1,397 mm SL female had a batch fecundity of 38,922,168 ±1,518,283 and 56,599,306 ±1,866,130 oocytes, respectively (Bullock and Smith 1991).
According to a conceptual model for the role of dispersal in a simple life history model of E. itajara, such groupers exhibit a positive response to the establishment of a marine reserve. Apart from inducing an increase in population growth rate, implementation of a reserve could increase population recovery rates by increasing reproductive output (Gerber et al. 2005).
Age, growth and longevity
Epinephelus itajara grow slowly relative to their potential maximum size. Growth rates for male and female are similar, averaging >100 mm per year through age 6, then slowing to about 30 mm per year by age 15, and finally declining to <10 mm per year after age 25. Von Bertalanffy growth model was found to be TL (mm) = 206[1-e(-0.126(Age+0.49)] (Bullock et al. 1992).
Maximum size and age recorded were 2,000-2,500 mm TL (Heemstra and Randall 1993), 37 years (female) and 26 years (male) (Bullock et al. 1992), respectively.
Habitat Type: Marine
Comments: Often found in shallow water; juveniles are common in mangrove swamps and both juveniles and adults occur in bays and harbors. Large adults are also encountered offshore on wrecks and in areas of high relief; they appear to occupy limited home ranges with little inter-reef movement (Heemstra and Randall 1993). Occurs from shallow bays and estuarine areas to offshore; large individuals have been observed in shallow waters of the Everglades, Florida Bay, and the Florida Keys (Bullock and Smith 1991). Inhabits shallow shore zones and is particularly prevalent near mangrove roots (Smith 1971, Thompson and Munro 1983). Habitat also includes shallow water, usually less than 30 m, but also artificial reefs and sunken reefs at 30-35 m depth (Colin 1990). Common nearshore around docks, in deep holes, wrecks, pilings, and cuts (Voss et al. 1969). Larger individuals only occur in places where big holes provide shelter (Nagelkerken 1981b). Characterized as an apex predator in shallow (< 30 m) reef and hard-bottom habitats (Parrish 1987). Juveniles occur in holes and below undercut ledges in swift tidal creeks draining mangroves; also occur in estuaries, especially around oyster bars, as well as mangrove channels (Tabb and Manning 1961, Bullock et al. 1992, Greenfield and Thomerson 1997). Center of abundance along Florida west coast appears to be the 10,000 Islands region near Naples due to extensive habitat (mangroves) for juveniles. Juveniles occur in mangrove areas where currents have undercut ledges or scoured out relatively deep holes (3 m) (Bullock and Smith 1991). Spawning aggregations have been observed during the summer on offshore wrecks at 36-46 m depth in the eastern Gulf of Mexico (Bullock and Smith 1991).
Water temperature and chemistry ranges based on 1 sample.
Depth range (m): 12.5 - 174
Temperature range (°C): 26.990 - 26.990
Nitrate (umol/L): 1.371 - 1.371
Salinity (PPS): 36.200 - 36.200
Oxygen (ml/l): 4.495 - 4.495
Phosphate (umol/l): 0.101 - 0.101
Silicate (umol/l): 3.321 - 3.321
Depth range (m): 12.5 - 174
Note: this information has not been validated. Check this *note*. Your feedback is most welcome.
Recorded at 100 meters.
Non-Migrant: No. All populations of this species make significant seasonal migrations.
Locally Migrant: No. No populations of this species make local extended movements (generally less than 200 km) at particular times of the year (e.g., to breeding or wintering grounds, to hibernation sites).
Locally Migrant: No. No populations of this species make annual migrations of over 200 km.
Diet: mobile benthic crustacea (shrimps/crabs), octopus/squid/cuttlefish, bony fishes, sharks/rays, sea snakes/mammals/turtles/birds
Comments: Smith (1971): eats mainly crustaceans.
Diseases and Parasites
Number of Occurrences
Note: For many non-migratory species, occurrences are roughly equivalent to populations.
Estimated Number of Occurrences: 6 - 20
Comments: Exact number of extant occurrences is unknown; may be limited to a few occurrences based upon known spawning aggregation sites in the eastern Gulf of Mexico (Colin 1990, Bullock and Smith 1991).
2500 - 10,000 individuals
Comments: Global abundance is unknown. Absent, disappearing, or becoming increasingly rare throughout range; especially true for larger individuals (Sobel 1996). The center of abundance is off peninsular Florida; more abundant in south Florida than in northern waters (Caribbean Fisheries Management Council 1993). Historically common in deeper channels in northern Florida Bay, Cape Sable; adults up to more than 400 pounds were taken in the Shark River estuary in the 1960s (Tabb and Manning 1961). Considered historically frequent in the Florida Keys (Starck 1968) but considered less common in the Florida Keys than in the Dry Tortugas (Jones and Thompson 1978). Commonly landed by Cuban Gulf Fleet operating off west Florida during the 1970s (Tashiro and Coleman 1977). Historically, the majority of the U.S. commercial catch was landed along the Gulf coast of Florida; landings were maximal in 1988, with 61,700 kg landed (Bullock et al. 1992). Prior to a moratorium on harvest, commercial landings in Monroe County, Florida, remained relatively stable between 1977 (14,558 kg) and 1989 (11,236 kg). Formerly very abundant in recreational landings; over 104,000 kg landed in 1982 compared to only 16,781 kg landed in 1989 (Bohnsack et al. 1994). Very rare in Aruba, Bonaire, and Curacao; uncommon in St. Martin, St. Eustatius, and Saba (Nagelkerken 1981a). Considered rare in Bermuda (Smith 1958). Also very rare in the central Bahamas; only one large individual (1.5 m) observed at 70 sites surveyed (Sluka et al. 1996).
Individuals tend to be site- or reef-specific, however, tagging studies have shown individuals may move 12 miles or more (Topp 1963).
Life History and Behavior
Transient spawning aggregations (Domeier and Colin 1997) have been observed from July-September in the eastern Gulf of Mexico (Colin 1990, Bullock and Smith 1991, Bullock et al. 1992). May pair-spawn like coney during August-September (Colin 1990). Spawning aggregations may consist of 20-30 large adults (45-200 kg) (Heemstra and Randall 1993). Batch fecundity for two females of 132 and 140 cm SL was 37-40 million and 55-58 million oocytes, respectively. Reproductive ability or maturation is not attained until a very large size: males first mature at 110-115 cm total length (4-6 years of age), while females first mature at 120-135 cm total length (6-7 years of age) (Bullock et al. 1992). Growth averages greater than 100 mm/year until the age of 6 years, when sexual maturity is attained; growth then declines to about 30 mm/year at age 15 (182-191 cm total length) and to less than 10 mm/year after age 25. Extremely long lived, with a lifespan of 30-50 years.
Molecular Biology and Genetics
Barcode data: Epinephelus itajara
Below is a sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species.
See the BOLD taxonomy browser for more complete information about this specimen and other sequences.
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Download FASTA File
Statistics of barcoding coverage: Epinephelus itajara
Public Records: 2
Specimens with Barcodes: 9
Species With Barcodes: 1
Statistics of barcoding coverage: Epinephelus quinquefasciatus
Public Records: 0
Specimens with Barcodes: 1
Species With Barcodes: 1
CITES: Not listed
IUCN Red List Assessment
Red List Category
Red List Criteria
Despite clear and promising signs of recovery in US waters following the 1990 moratorium, the increases in numbers noted are young and juvenile fish (the species takes five to six years to become sexually mature). Hence many years of protection are still needed to enable populations to recover reproductive potential and range. Continued surveys and education programmes for this species and its inclusion in marine protected areas are proposed.
Because of the factors listed above, this species is listed as Critically Endangered (A2d).
It is recommended that the species be reassessed after five years following the completion of dedicated surveys and stock assessments.
National NatureServe Conservation Status
Rounded National Status Rank: NNR - Unranked
NatureServe Conservation Status
Rounded Global Status Rank: G2 - Imperiled
Reasons: Relatively wide geographic range in tropical and subtropical waters of the Atlantic and eastern Pacific oceans; greatest abundance occurs off peninsular Florida; significant population declines due to overfishing seem to have occurred throughout range, but principally in the eastern Gulf of Mexico and southeastern U.S.; considered extremely vulnerable to fishing because of life history characteristics; there is little, if any, evidence of recovery of overfished populations; status of populations in the eastern Pacific is unknown.
Other Considerations: Vulnerable to excessive exploitation due to slow growth, delayed reproduction, and aggregate spawning.
Female-to-male sex ratio was 1.75:1 in the eastern Gulf of Mexico from 1977 to 1990 (Bullock et al. 1992).
Distribution densities of juveniles around mangrove islands (22-61 juveniles per km of mangrove shoreline) in the Ten Thousand Islands were found to be higher and less variable than the densities in rivers (0-46 juveniles per km) (Koenig et al. 2007).
Sexual pattern has not been confirmed (Bullock et al. 1992, Sadovy and Eklund 1999).
Most individuals collected from the eastern Gulf of Mexico were between nine and 15 years (with female and male ages overlapping), few exceeded 30 years (Bullock et al. 1992).
Estimated average length and fishing mortality in the exploited stock were found to be 1,161 mm total length (TL) and 0.04 per year in the Florida Keys, respectively (Ault et al. 2005).
Based on the distribution of age classes (Bullock et al. 1992) and a maturity age of six years (Sadovy and Eklund 1999), the generation time for Atlantic Goliath Grouper is estimated at 13.5 years (Note: Generation time is defined here as the mean age of reproductive individuals in a population). Therefore, measures of declines and/or recovery over three generations covers a time span of 40.5 years.
Global Short Term Trend: Decline of 30 to >90%
Comments: Most severely impacted grouper in the western Atlantic; overfishing led to severe reductions in populations (Sadovy 1990). Presently scarce throughout the Gulf of Mexico, U.S. western Atlantic, Puerto Rico, and the U.S. Virgin Islands; no longer abundant in any part of range (Caribbean Fisheries Management Council 1993). Declines in landings led to fishery closures in the state of Florida and U.S. territorial waters in 1990 (Bohnsack et al. 1994). Considered overfished in the Florida Keys based on low spawning potential ratio (Ault et al. 1998). Not common in the Netherlands Antilles (Nagelkerken 1981b). Specimens over 500 pounds were frequently observed while diving offshore of Tampa Bay and other areas of the eastern Gulf of Mexico in the 1950s and 1960s (Springer and Woodburn 1960).
Epinephelus itajara is apparently vulnerable to stresses caused by cold water (Gilmore et al. 1978) or red tide — it was recorded that populations of E. itajara were much reduced during a red tide in 1971 and dead individuals over 45 kg were often observed (Smith 1976). During an outbreak of red tide in Florida in March 2003, eleven large dead Goliath Grouper (sized 305 mm to 2,057 mm) washed up near the Sanibel Island Causeway (www.sefsc.noaa.gov/redtidegrouper.jsp, accessed on 4th Jan 2006).
Degree of Threat: A : Very threatened throughout its range communities directly exploited or their composition and structure irreversibly threatened by man-made forces, including exotic species
Comments: Historically this species was of commercial and recreational importance in the western central Atlantic, but now it is considered the most impacted grouper species in the region (Sadovy 1990). Considered very threatened range-wide, principally from fishing and life history characteristics: slow growth, longevity, low natural mortality, and high vulnerability during spawning aggregations (Bullock et al. 1992, Heemstra and Randall 1993). Fishermen historically exploited spawning aggregations in the Key West area.Considered overfished in the Florida Keys based upon a spawning potential ratio less than 30 percent (Ault et al. 1998). Of minor importance in commercial fisheries off the west coast of Florida, where landings for 1988 totaled 61,700 kg; easily approached and speared by divers, hence they are scarce in areas accessible to divers (Bullock et al. 1992). Caught with hook-and-line, occasionally in traps and trawls. Chief method of harvest in Puerto Rico and the U.S. Virgin Islands is spear fishing (Caribbean Fisheries Management Council 1993). Presently protected in U.S. territorial waters and the U.S. Virgin Islands.
The NMFS, under the authority of the Magnuson-Stevens Fisheries Conservation and Management Act, lists the E. itajara as overfished in Reports to Congress on the Status of Fisheries (www.sefsc.noaa.gov/redtidegrouper.jsp accessed on 4th Jan 2006, Porch et al. 2003).
The American Fisheries Society classified E. itajara as being conservation dependent. Due to the fish's life history, it is vulnerable to become threatened, but can be kept from threatened status with appropriate protective measures (www.gulfcouncil.org/oldstories/2000-07-20-jewfish-update.htm, accessed on 31st Dec 2005).
Classified as endangered by US Fish and Wildlife Service (USFWS) under the concept of District Population Segments (DPS) but regarded as recovering under Fisheries Management Plan (FMP) (Musick et al. 2000).
A 5-year-protection was granted by the IBAMA (Brazilian Environmental Agency) on 20th September 2002 (www.vidamar.org.br/meros/english/index.php accessed on 3rd Jan 2006).
According to the regulations for E. itajara fishing in Puerto Rico and the US Virgin Islands, fishing in any form for this species is totally prohibited all year around (www.caribbeanfmc.com; accessed on 4th Jan 2006).
It was indicated that the ban on spear-fishing in the upper Florida Keys has significantly and beneficially influenced the average size of groupers, although their populations in this region have not reached stable levels (Sluka and Sullivan 1998).
Biological Research Needs: More research on population dynamics and reproductive biology is needed. A greater understanding of reproductive biology would greatly facilitate stock management. There is a need to know (1) whether most annual reproduction occurs at spawning aggregations and (2) the geographic locations and durations of significant aggregations. There is also a need to know how aggregation fishing is likely to affect courtship or spawning behavior. Relatively little is known concerning the minimum size of sexual maturation relative to the size of entry into the fishery. It is also not known to what extent individuals recruit locally or from larvae from up-current and/or off-island locations. Little is known concerning the characteristics of critical juvenile habitat nor of the principal settlement periods (Sadovy 1990).
Global Protection: Few to several (1-12) occurrences appropriately protected and managed
Comments: The South Atlantic Fishery Management Council administers commercial and recreational fisheries in U.S. territorial waters of the Atlantic. Federal commercial regulations prohibit the possession or harvest of jewfish in or from the Economic Exclusive Zone. Prior to the harvest ban in 1990, a minimum size limit of 13 inches (fork length) was established in Florida waters before 1970. In 1985, the size limit was increased to 18 inches (Huntsman et al. 1990). Harvest ban in U.S. federal waters and state waters of Florida became effective in 1990 because of vulnerability to overfishing and dramatic declines in populations (Bullock et al. 1992, Bohnsack et al. 1994). Because it is rare and easily taken, it is now offered stringent protection in Texas, Louisiana, and Florida, where possession by anglers is prohibited (Hoese and Moore 1998). It is unlawful to harvest, possess, land, purchase, sell or exchange the species in Florida. Also protected from collection in the U.S. Virgin Islands (Caribbean Fisheries Management Council 1993).
Needs: Occurrences should be protected to ensure long-term survival. Stocks should be managed to maintain adequate reproductive (spawning) stock biomass to sustain recruitment. Serious consideration should be given to the protection of spawning aggregations, at a minimum by prohibiting spear guns and fish traps and by permitting only low levels of exploitation at most. The introduction of no-fishing zones in critical areas also needs to be considered as a management option to address the problem of overfishing (Sadovy, in press).
Relevance to Humans and Ecosystems
Comments: Caribbean Fisheries Management Council (1993): chief method of harvest is spear-fishing in Puerto Rico and the U.S. Virgin Islands.
Atlantic goliath grouper
The Atlantic goliath grouper or itajara (Epinephelus itajara), also known as the jewfish, is a large saltwater fish of the grouper family found primarily in shallow tropical waters among coral and artificial reefs at depths from 5 to 50 m (16 to 164 ft). Its range includes the Florida Keys, the Bahamas, most of the Caribbean, and practically all of the Brazilian coast. On some occasions, it is caught in New England off Maine and Massachusetts. In the eastern Atlantic Ocean, it occurs from the Congo to Senegal.
They may reach extremely large sizes, growing to lengths of up to 2.5 m (8.2 ft) and can weigh as much as 360 kg (790 lb). The world record for a hook and line-captured specimen is 309 kg (681 lb), caught off Fernandina Beach, Florida, in 1961. They are usually around 180 kg (400 lb) when mature. Considered of fine food quality, Atlantic goliath grouper were a highly sought-after quarry for fishermen. The grouper's inquisitive and generally fearless nature makes it a relatively easy prey for spear fishermen. They also tend to spawn in large aggregations, returning like clockwork to the same locations, making them particularly vulnerable to mass harvesting.
Until a harvest ban was placed on the species, its population was in rapid decline. The fish is entirely protected from harvest and is recognized as a critically endangered species by the IUCN. The US began protection in 1990, and the Caribbean in 1993. The species' population has been recovering since the ban; with the fish's slow growth rate, however, it will take some time for populations to return to their previous levels.
Goliath groupers are believed to be protogynous hermaphrodites, which refer to organisms that are born female and at some point in their lifespan change sex to male. Most grouper follow this pattern, but it has not yet been verified for the goliath. In fact, males could be sexually mature at smaller sizes (~1150 mm) and younger ages (4–6 years) than females (~1225 mm and ~6–8 years).
The Atlantic goliath grouper has been referred to as the jewfish, but in 2001 the American Fisheries Society stopped using that term because they felt that it was "culturally insensitive".
- Craig, M.T. (2011). "Epinephelus itajara". IUCN Red List of Threatened Species. Version 2013.2. International Union for Conservation of Nature. Retrieved 30 March 2014. Database entry includes a range map and a lengthy justification of why this species is critically endangered
- IGFA World Records
- "FLMNH Ichthyology Department: Goliath Grouper". Flmnh.ufl.edu. Retrieved 2011-10-31.
- Bullock et al. (1992). Age, Growth, and Reproduction of Jewfish Epinephelus itajara in the Eastern Gulf of Mexico. U.S. Fishery Bulletin 90 (2):243-249. Retrieved August 21, 2014.
- Brassfield, Mike (May 24, 2001). "Big fish get a giant name". St. Petersburg Times. Retrieved August 21, 2014.
Names and Taxonomy
Comments: Also known as mero (Brazil), mero guasa (Colombia), cherne (Peru), guasa (Cuba, Venezuela), and (formerly) jewfish (U.S.).
Smith (1971): considered part of subgenus Promicrops, which was previously considered a genus. Appears to be related to Epinephelus lanceolatus and E. tauvina of the Indo-Pacific. The subgenus is demarcated by a broading and depressing of the head and body. No close relatives in American waters.
DNA data indicate that (1) American Epinephelus species are not monophyletic, (2) Cephalopholis is monophyletic only with the inclusion of the morphologically distinct Paranthias, and (3) Mycteroperca is monophyletic only with the inclusion of the Indo-Pacific Anyperodon leucogrammicus (Craig et al. 2001).
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