Description and Distribution
Red hind (Epinephelus guttatus) is one of the more common groupers on coral reef habitats whose distribution ranges from Brazil to Bermuda. It occurs in Bermuda, throughout the Bahamas, in the greater and lesser Antilles, and along the Central and South American coasts. The species is important in terms of total landings in the Caribbean and provides excellent fish meat. It is easily approached by divers and is readily caught on hook and line and speared. It feeds mainly on crabs and other crustaceans, fishes, and octopus. (24, 28)
E. guttatus occupies shallow coral reef and rocky bottom from 3 to 50 m depth (28).
The above diagram shows the range of red hind (highlighted in green)
and reported spawning aggregation sites (highlighted in red)
Red hind is usually solitary but seasonally aggregates to spawn. It is a protogynous hermaphrodite which changes sex from female to male at 28 to 38 cm (TL). It normally reaches 50 to 55 cm (TL) and 11 years. Maximum reported length is 72 cm (TL) and age is 22 years. Most fish larger than 40 cm (TL) are males. It produces pelagic eggs and larvae. Eggs hatch in 27 hours in the laboratory. Females rest on or close to the bottom, while males patrol around an area that consists of one to five female(s), and defend this territory from other males. Red hinds form aggregations and reproduce almost exclusively within the aggregation period which occurs from December to April in the Caribbean, and from May to July in Bermuda. Spawning aggregations of red hinds have been studied in a number of locations throughout the Caribbean, including Bermuda and Saba, the British Virgin Islands, Jamaica, Netherland Antilles, Puerto Rico, and the United States Virgin Islands (USVI) (6,7,10,12,13,16,18,23,26,27,28,30,31,32,33).
Sexual pattern and spawning aggregations
E. guttatus females are determinate spawners and spawn more than once during the course of the several months long annual spawning season. Spawning aggregations typically occur on the top of deep coral reef ridges located on or near the shelf edge, 1 to 2 m above the reef. In Jamaica, Puerto Rico and USVI, the aggregations form at the week before full moon from December to February. Spawning typically peaks in January and aggregations completely disperse following the January or February full moon. In Bermuda, the species aggregate to spawn also at full moon but the months differ from elsewhere; May, June and July. Red hinds migrate 5 to 33 km from home sites to spawning grounds and exhibit a functional spawning migration area of 90 to 500 km2. Swim speeds of 3 km d-1 over 20 km during migration have been recorded for red hinds in Bermuda.
Spawning typically occurs during periods of declining seawater temperature and slacking currents within a temperature range of 26º to 27.5º. Males arrive several weeks before spawning at spawning site and stay for the duration of the spawning season whereas females may only spawn during one month, sometimes longer. Gender-specific movements during the week of spawning cause daily fluctuations in female:male sex ratios that range from more than 20:1 to less than 1:1. In between monthly spawning peaks, about 5-20 % of the spawning population remain on the aggregation site (2,5,10,13,18,19,27,29,30).
Hinds and groupers (Serranidae) form a valuable component of reef fisheries throughout the Caribbean and tropical western Atlantic. Red hind, in particular, contributed 67% to 99% of the total annual commercial fin fish catch in the USVI between 1987 and 1991, and over 30% of the grouper catch in Bermuda. In Puerto Rico, red hind is the most important grouper, following the decline of E. striatus. Fishers have exploited spawning aggregations of the species for years, leading to adverse consequences to populations (1,2,7,12,15,16,25).
No information. There is no reported research or commercial culture of the species.
E. guttatus is listed as Least Concern on the IUCN red list (assessed 2008) as it is a currently widespread and abundant species, albeit heavily fished. It may be of conservation concern in the future, especially if its aggregations are not protected effectively (28).
In the Caribbean, the species is threatened by the destruction of coastal habitats. Commercial and recreational fisheries target red hinds with SCUBA with speargun, hook and line, fish traps and nets. Much effort towards red hind is expanded during the reproductive season of the species despite local regulations. Other fishing gear such as traps and nets that do not target red hind may take them incidentally (6,7,28).
Also, males of several grouper species, including red hind, remain higher in the water column than females defending spawning territories or displaying to potential mates. Sex and/or size -selective fishing mortality at spawning aggregations may occur when these more behaviourally dominant males take baited hook more frequently than females (5,8,11).
Fishery-independent data trends in Puerto Rico showed annual progressive decreases in average catch per unit effort (CPUE) and average lengths of males and females in both protected and unprotected areas, prior to the enactment of seasonal closures (in 2004, see below). A small increase in annual average CPUE occurred in the region soon after the enactment of closures, but was followed by continual low values. Initially, there were no significant differences in annual average lengths, by sex, prior to and following enactment of seasonal closures, but significant increases were observed during 2004 to 2006 (12,28).
Conservation & Management
In the Caribbean, different management measures have been used to protect red hind. It is currently present in coral reef protected areas in the Greater Caribbean and Florida and regulated by fisheries department in the region. These include permanent closures (St. Thomas, Bermuda), seasonal closures (St. Croix), and seasonal market closures (British Virgin Islands, western Puerto Rico). The effectiveness of some of these protected areas is not well-known (18,20).
There is not enough population information on the effects of spatial closures in sustaining red hind spawning aggregations. However, several studies in the USVI indicate that seasonal closures may improve characteristics (length, sex ratio) of some aggregations but not others, whereas large permanent closures can facilitate rapid recovery of a spawning population (18,20).
In 2004, Puerto Rico's Department of Natural Resources passed a fisheries regulation to prohibit the take of E. guttatus during its spawning season (December 1 to February 28 of each year), throughout the waters of Puerto Rico (up to 16.7 km from shore). This eliminated the site-specific protection measures of known spawning areas that had already been in place for some time. Outside the 16.7km (exclusive economic zone) the government of the United States (National Oceanic and Atmospheric Administration) designated a ban on the take of red hind but enforcement and implementation of the new regulations has been low (28).
Spawning aggregations of red hind are protected in St. Thomas (US Virgin Islands), and there are signs that the species is recovering in this area, including an increase in average size of fishes and better catch rates outside of the aggregation period (18).
1. Beets J, Friedlander A (1992) Stock analysis and management strategies for red hind, Epinephelus guttatus, in the U.S. Virgin Islands, Vol 42. 42nd Gulf and Caribbean Fisheries Institute:66-79
2. Beets J, Friedlander A (1999) Evaluation of a conservation strategy: a spawning aggregation closure for red hind, Epinephelus guttatus, in the U.S. Virgin Islands. Environmental Biology of Fishes 55:91-98
3. Coleman FC, Koenig CC, Collins LA (1996) Reproductive styles of shallow-water groupers (Pisces: Serranidae) in the eastern Gulf of Mexico and the consequences of fishing spawning aggregations. Environmental biology of fishes 47:129-141
4. Colin PL (1992) Reproduction of the Nassau grouper, Epinephelus striatus, (Pisces: Serranidae) and its relationship to environmental conditions. Environmental Biology of Fishes 34:357-377
5. Colin PL, Shapiro DY, Weiler D (1987) Aspects of the reproduction of two groupers, Epinephelus guttatus and E. straitus in the West Indies. Bulletin of Marine Sciences 40:220-230
6. Cummings NJ, Parrack ML, Zweifel JW (1997) The status of red hind and coney in the U.S. Virgin Islands between 1974 and 1992, Vol 49. 49th Gulf and Caribbean Fisheries Institute:354-397
7. Eristhee N, Kadison E, Murray PA, Llewelyn A (in press) Preliminary investigations into the red hind fishery in the British Virgin Islands, Vol. 57th Gulf and Caribbean Fisheries Institute St. Petersburg, FL
8. Gilmore RG, Jones RS (1992) Colour variation and associated behaviour in the Epinepheline groupers, Myceroperca microlepis (Goode and Bam) and M. phenax (Jordan and Swain). Bulletin of Marine Sciences 51:84-103
9. Huntsman GR, Potts J, Mays RW, Vaughan DS (1999) Groupers (Serranidae, Epinephelinae): Endangered apex predators of reef communities. In: Musick JA (ed) Life in the slow lane. Ecology and Conservation of long-lived marine animals. American Fisheries Society, Bethesda, p 217-232
10. Kadison E, Nemeth RS, Blondeau JE (in press) Investigation of a red hind (Epinephelus guttatus) spawning aggregation on Saba Bank in Netherlands Antilles. Bulletin of Marine Sciences
11. Levin PS, Grimes CB (2002) Reef fish ecology and grouper conservation and management. In: Sale PF (ed) Coral reef fishes. Dynamics and diversity in a complex ecosystem. Academic Press, London, p 377-389
12. Luckhurst BE (1996) Trends in commercial fishery landings of groupers and snappers in Bermuda from 1975 to 1992 and associated fishery management issues. In: Arreguin-Sanchez F, Munro JL, Balgos MC, Pauly D (eds) Biology, fisheries and culture of tropical groupers and snappers, Vol 48. ICLARM, p 286-297
13. Luckhurst BE (1998) Site fidelity and return migration of tagged red hinds (Epinephelus guttatus) to a spawning aggregation site in Bermuda. Proceedings of 50th Gulf and Caribbean Fishery Institutes 50:750-763
14. Luckhurst BE, Barnes J, Sadovy Y (1992) Record of an unusually large red hind, Epinephelus guttatus (Serranidae), from Bermuda with comments on its age. Bulletin of Marine Sciences 51:267-270
15. Matos-Caraballo D. (2000) Portrait of the commercial fishery of the red hind Epinephelus guttatus in Puerto Rico during 1992-99. Gulf and Caribbean fisheries institute 53th annual meeting. 16 pp.
16. Munro JL, Blok L (2003) The Status of stocks of groupers and hinds in the Northeastern Caribbean. Proceedings of 56th Gulf and Caribbean fisheries institute annual meeting 56:283-294
17. Munro JL, Gout VC, Thompson R, Reeson P (1973) The spawning season of Caribbean reef fishes. Journal of Fish Biology 5:69-84
18. Nemeth RS (2005) Population characteristics of a recovering US Virgin Islands red hind spawning aggregation following protection. Marine Ecology Progress Series 286:81-97
19. Nemeth RS, Blondeau J, Herzlieb S, Kadison E (2007) Spatial and temporal patterns of movement and migration at spawning aggregations of red hind, Epinephelus guttatus, in the U.S.Virgin Islands. Environmental Biology of Fishes 78:365-381
20. Nemeth RS, Herzlieb S, Blondeau J (2006) Comparison of two seasonal closures for protecting red hind spawning aggregations in the US Virgin Islands. Proceedings of 10th International Coral Reef Conference 4:1306-1313
21. NMFS (1999) Our living oceans. Report on the status of U.S. living marine resources, 1999. Report No. NMFS-F/SPO-4, Department of Commerce, Washington, D.C.
22. Olsen DA, LaPlace JA (1978) A study of Virgin Islands grouper fishery based on a breeding aggregation. Proceedings of 31st Gulf and Caribbean Fishery Institute 31:130-144
23. Randall JE (1983) Caribbean Reef Fishes, Vol. TFH Publications, Neptune City, NJ
24. Robins CR, Ray GC (1986) A field guide to Atlantic coast fishes, Vol. Houghton Mifflin Company, Boston.
25. Sadovy Y, Figuerola M (1992) The status of red hind fishery in Puerto Rico and St. Thomas, as determined by yield-per-recruit analysis. Proceedings of 42nd Gulf and Caribbean Fishery Institute 42:23-38
26. Sadovy Y, Figuerola M, Roman A (1992) Age and growth of red hind Epinephelus guttatus in Puerto Rico and St. Thomas. Fisheries Bulletin 90:516-528
27. Sadovy Y, Rosario A, Roman A (1994) Reproduction in an aggregating grouper, the red hind, Epinephelus guttatus. Environmental Biology of Fishes 41:269-286
28. Sadovy, Y., Rocha, L., Choat, J.H., Bertoncini, A.A., Ferreira, B.P. & Craig, M. 2008. Epinephelus guttatus. In: IUCN 2008. 2008 IUCN Red List of Threatened Species. <www.iucnredlist.org>
29. Shapiro DY (1987) Reproduction in groupers. In: Polovina JJ, Ralston S (eds) Tropical snappers and groupers: biology and fisheries management. Westview Press, Boulder, p 295-327
30. Shapiro DY, Sadovy Y, McGehee MA (1993) Size, composition, and spatial structure of the annual spawning aggregation of the red hind, Epinephelus guttatus (Pisces: Serranidae). Copeia 2:399-406
31. Smith CL (1971) A revision of the American groupers: Epinephelus and allied genera. Bulletin of the American Museum of Natural History 146:69-241
32. Thompson R, Munro JL (1978) Aspects of the biology and ecology of Caribbean reef fishes: Serranidae (hinds and groupers). Journal of Fish Biology 12:115-146
33. Whiteman EA, Jennings CA, Nemeth RS (2005) Sex structure and potential female fecundity in a red hind (Epinephelus guttatus) spawning aggregation: applying ultrasonic imaging. Journal of Fish Biology 66:983-995