REFE!2EPJ65T;: Do Not Remove fr2m the Library U. S. and Wildlife Service National Weilcnds Resmrch Center Blologlcal Report 82(11.110) /00 Cajun Dome Boulevard TR EL-824 ~ugust1989 Lafayette, Louisiana 70506

Species Profiles: Life Histories and Environmental Requirements of Coastal and Invertebrates (South )

BLACK, RED, AND NASSAU

Coastal Ecology Group Fish and Wildlife Service Waterways Experiment Station U.S. Department of the Interior U.S. Army Corps of Engineers Biological Report 82(11.110) TR EL-82-4 August 1989

Species Profiles: Life Histories and Environmental Requirements of Coastal Fishes and Invertebrates (South Florida)

BLACK, RED, AND NASSAU GROUPERS

Darryl E. Jory and Edwin S. Iversen University of Miami Rosenstiel School of Marine and Atmospheric Science Division of Biology and Living Resources 4600 Rickenbac ker Causeway Miami, FL 33149

Project Officer David Moran U.S. Fish and Wildlife Service National Wetlands Research Center 1010 Gause Boulevard Slidell, LA 70458

Performed for Coastal Ecology Group U. S. Army Corps of Engineers Waterways Experiment Station Vicksburg, MS 39180

and

U.S. Department of the Interior Fish and Wildlife Service Research and Development National Wetlands Research Center Washington, DC 20240 DISCLAIMER

The mention of trade names in this report does not constitute endorse- ment nor recommendation for use by the U.S. Fish and Wildlife Service or the Federal Government.

This series may be referenced as follows: U.S. Fish and Wildlife Service. 1983-19-. Species profiles: life histories and environmental requirements of coastal fishes and invertebrates. U.S. Fish Wildl. Serv. Biol. Rep. 82(11). U.S. Army Corps of Engineers, TR EL-82-4.

This profile may be cited as follows: Jory, D.E., and E.S. Iversen. 1989. Species profiles: life histories and environmental requirements of coastal fishes and invertebrates (south Florida) --black, red, and Nassau groupers. U.S. Fish Wi ldl . Serv. Biol . Rep. 82(11.110). U.S. Army Corps of Engineers, TR EL-82-4. 21 pp. PREFACE

This species profile is one of a series on coastal aquatic organisms, principally fish, of sport, commercial, or ecological importance. The profiles are designed to provide coastal managers, engineers, and biologists with a brief comprehensive sketch of the bio1ogical characteristics and environmental requirements of the species and to describe how populations of the species may be expected to react to environmental changes caused by coastal development. Each profile has sections on , 1ife history, ecological role, environmental requirements , and economic importance, if appl icable. A three-ring binder is used for this series so that new profiles can be added as they are prepared. This project is jointly planned and financed by the U. S. Army Corps of Engineers and the U.S. Fish and Wildlife Service.

Suggestions or questions regarding this report should be directed to one of the following addresses.

Information Transfer Specialist National Wetlands Research Center U.S. Fish and Wildlife Service NASA-Slidell Computer Complex 1010 Gause Boulevard Slidell, LA 70458

U. S. Army Engineer Waterways Experiment Station Attention: WESER-C Post Office Box 631 Vicksburg, MS 39180 CONVERSION TABLE

Metrlc to U.S. Customary

Multlply To Obtain millimeters (mm) inches centimeters (cm) inches meters (m) feet meters (m) fathoms kilometers (km) statute miles kilometers (km) nautical miles square meters (m2) square feet square kilometers (km2) square miles hectares (ha) acres liters (I) gallons cubic meters (m3) cubic feet cubic meters (m3) acre-feet milligrams (mg) ounces grams (g) ounces kilograms (kg) pounds metric tons (t) pounds metric tons (t) short tons kilocalories (kcal) British thermal units Celsius degrees (OC) Fahrenheit degrees U.S. Customary to Metric inches millimeters inches centimeters feet (ft) meters fathoms meters statute miles (mi) kilometers nautical miles (nmi) kilometers square feet (ft2) square meters square miles (mi2) square kilometers acres hectares gallons (gal) liters cubic feet (ft3) cubic meters acre-feet cubic meters ounces (02) milligrams ounces (02) grams pounds (Ib) kilograms pounds (Ib) metric tons short tons (ton) metric tons

British thermal units Ptu) kilocalories Fahrenheit degrees ( F) Celsius degrees CONTENTS

Page PREFACE ...... iii CONVERSION TABLE ...... iv ACKNOWLEDGMENTS ...... vi NOMENCLATLIRE/TAXONOMY ...... 1 REASONS FOR INCLUSION IN THE SERIES ...... 1 GEOGRAPHIC RANGE ...... 2 MORPHOLOGY AND IDENTIFICATION AIDS ...... 2 Morphological Characteristics ...... 2 Morphological Differences ...... 4 Color and Pigmentation ...... 5 Size ...... 5 LIFE HISTORY ...... b Habitat...... 6 Sex Reversal and Spawning ...... 6 Eggs and Larvae ...... 7 Juveniles ...... 8 Movement and Migration...... 8 GROWTH CHARACTERISTICS ...... 9 FISHERIES ...... 9 Recreational and Commercial Fisheries...... 9 Marketing ...... 10 Regulations ...... 10 Artificial Reefs ...... 10 Contaminants ...... 11 ECOLOGICAL ROLE ...... 11 Food and Feeding Habits ...... 11 Competitors and Predators ...... 13 Parasites and Diseases ...... 13 Ciguatera (Tropical Fish Poisoning) ...... 15 Behavior ...... 15 REFERENCES ...... 17 ACKNOWLEDGMENTS

We are grateful for the useful references contributed by C. S. Manooch 111 of the National Marine Fisheries Service, and for his thorough review. S. Bortone of the University of West Florida also reviewed the manuscript. E. Snell and R. Vaught of the National Marine Fisheries Service provided catch data and pertinent literature respectively. B.E. Luckhurst also made available useful literature. J. Iversen reviewed the general draft and made numerous useful comments. G. A. Maury prepared the cover and Figure 1. V. R. Kestrepo, E. Lahman, J.F. Carranza, and I. Paez kindly assisted with word processing and greatly facilitated the completion of this report. r

BLACK

0- 100 Mn

Figure 1. Black, red, and Nassau groupers.

BLACK, RED, AND NASSAU GROUPERS

NOMENCLATURE/TAXONOMY REASONS FOR INCLUSION IN THE SERIES Scientific name ...... Black, red, and Nassau groupers bonaci Poey, 1860 are actively sought by both commercial Preferred common name....Black and sport fishermen throughout their Other common names...Marbled rockfish, geographic ranges. The red grouper is black rockfish, snider grouper, car- one of the most abundant of the 17 barita, junef ish (Figure 1) species of grouper caught in Florida by commerci a1 and recreational f ishermen. The 1984 grouper landings for Florida Scientific name ...... totaled about 10 million lb, worth $14 morio Valenciennes, 1828 million. The south Florida region Preferred common name...... Red grouper (from Citrus around to Brevard Coun- (Figure 1) ties) contributed more than 8 million lb (81% of the total, worth more than $11 million (about 79% of the total). Scientific name ...... Epinephelus The 1984 Florida finfish landings were striatus Bloch, 1792 worth about $57.5 million, of which Preferred common name...Nassau grouper groupers contributed $14 mi11 ion or Other common names ...... Grouper, nearly 25% of the total--making them rockfish, hamlet (Figure 1) the most valuable marine finfish group in Florida. However, despite their economic importance, available infor- Class ...... Osteichthyes mation on their life histories in the Order ...... south Florida region--particularly Family ...... for black and Nassau groupers--is grossly inadequate for effective man- Chapl in 1968; Smith 1971; Fi scher agement of the fishery. Beaumar iage 19781, although the three species are and Bullock (1976) wrote that fewer rare north of Florida. Reports of than two dozen pertinent studies had these species north of the Carolinas then been published on the biology of are probably a result of larval groupers, and that almost half of transport by currents, as suggested by these were based on tagging programs Thompson and Munro (1978) for other that described movement patterns. Ac- grouper species. cording to these authors "This paucity of data exemplifies the need for simi- The black grouper is abundant in lar life histories studies, if sound the Florida Keys (Randall 1968), in management policies are to be adopt- , and off and ed. ..." Venezuela (Cervigon 1966); it is re- portedly rare in the eastern Gulf of These three species of groupers Mexico (Smith et al. 1975), the Virgin are an important component of the Islands, Puerto Rico, and the highly diverse reef fish community, Colombian Caribbean (Dahl 1971). occupying positions near the top of the food webs. Their biology, ecology, The red grouper is primarily a and exploitation are integral compo- continental species, having the widest nents of fisheries and distribution of all western central management of reef fisheries (Bohnsack Atlantic groupers (Roe 1976). It is 1982; Bannerot 1984). Any significant found mostly in broad shelf areas; its change in their numbers could affect center of abundance is in the Florida the balance of reef ecosystems (May et shelf and the eastern al. 1979). Preservation of coral reefs (Moe 1969). It is also abundant in the is essential for the perpetuation of Colombian Caribbean (Dah 1 1971), and harvestable grouper populations. off northeastern Venezuela (Cerv igon 1966), but uncommon in the In south Florida, ciguatera (poi- (Randal 1 1968). soning from consumption of tropical fishes) is endemic. Many of the The Nassau grouper is primarily reported cases from black grouper are an insular species, very common in really from incorrectly identified fish the West Indies (Randall 1968), the smuggled in from the Bahamas, where Bahamas (Boh 1ke and Chapl in 1968), ciguatera is much more common (Jones southern Gulf of Mexico (Fischer Bohnsack, National Marine Fisheries 1978), and the Colombian Caribbean Service, Miami, FL; pers. comm.). Life (Dahl 1971). In Venezuela, it is com- history data, especially on feeding mon in the Archipelago Los Roques but habits, of the species implicated are rare in northeastern islands such as needed to help prevent ciguatera. The Margarita, Coche, and Cubagua toxin causes gastrointestinal, cardio- (Cervigon 1966). Red and Nassau grou- vascular, and neurological disturbances pers occur sympatrically in the Flori- resulting in pro1onged disabi 1i ty and da Keys (Figure 2), although their 1ong and expensive recovery periods local distribution is essentially dis- (de Sylva and Higman 1980; Poli 1982). junct (Moe 1969).

GEOGRAPHIC RANGE MORPHOLOGY AND IDENTIFICATION AIDS

The black, red, and Nassau grou- pers range from New England and Morphological Characteristics to southeastern Brazi 1, including the Bahamas, Gulf of Mexico Smith (1971) provided the fol- and all of the Caribbean (B6hlke and lowing morphological descriptions. T. PETERSBURG

FLORIDA

HOUSAND ISLANDS

MILES

Area of distribution p

Figure 2. Approximate distribution of the black, red, and Nassau groupers in south Florida. 3 Black grouper. "Large species of and only a slightly emarginated notch, Mycteroperca with robust body, rela- and from other serranids in having the tively large scales, and rounded bases of the soft dorsal and anal fins preopercle. Dorsal fin XI, 17; anal covered by scales and thick skin. fin 111, 12(13); pectoral fins 17; gill rakers 20-26. Posterior nostril Red grouper. This grouper can be not enlarged. Vertical fins without distinguished from other species of exserted rays. First three dorsal the genus Epinephelus by its dorsal spines low, not forming an elevated fin, in which the second spine is the lobe. Gill rakers are moderate in longest and the interspinous membrane length. The exposed surface of the is not notched. InE. nigritus, E. maxilla is scaled. The upper lip is f lavol imbatus, E. mystacinus, and E. broad, as wide as or wider than the niveatus, the pelvic fins are longer shaft of the maxilla at the midpoint than the pectoral fins and are inser- of the supramaxi 11ary groove. " ted anterior to the pectoral-fin base, whereas in the red grouper the pelvic Red grouper. "Moderate-sized spe- fins are shorter than the pectorals cies of Epinephelus with large eyes, and are inserted slightly behind the small scales, robust body. Nostrils ventral end of the pectoral fin base. subequal, posterior little larger than The red grouper differs from anterior. Dorsal fin XI, 16-17 with Mycteroperca species in having a less interspinous membrane not notched. elongated body, and in having nine Second dorsal spine longest. Vertical soft anal-fin rays. The red grouper fins angulate in large fish. Anal fin differs from other serranids in 111, 9; pectoral fins 17; gill rakers having a more robust body, having the 23-25. " bases of tbe dorsal and anal fins covered by scales and thick skin, and Nassau grouper. "Moderate-sized in having 11 dorsal fin spines. species of Epinephelus with large eyes, medium-sized scales, robust Nassau grouper. This grouper can be body. Nostri 1s subequal, -posterior distinauished from other species in sl ight ly enlarged, comma-shaped in the g&us Epinephelus by the third adults. Dorsal fin XI, 16-17 with spine of the dorsal fin which is interspinous membranes notched; anal longer than the second, and in having fin modally 111, 8; pectoral fins 18; a slightly indented interspinous mem- gill rakers 24-25. Vertical fins brane. The caudal fin is slightly rounded. " emarginated. In Nassau groupers the pelvic fins are shorter than the pec- torals and are inserted below or Morpholoqical Differences behind the ventral end of the pectoral fin base. In E. mystacinus, -E. Morphological differences among nigritus, -E. flavolimbatus, and -E. grouper species were cited by Fischer niveatus. the ~elvicfins are lonaer (1978). than the' pectdrals and are inserted anterior to the ventral end of the Black grouper. This grouper can be base of the pectoral fins. In addi- distinguished from species of the tion, E. mystacinus has greatly en- genus Epinephelus by its more elon- 1arged, equal-sized posterior nos- gated body, which is not deepest at tri 1s, whereas in the Nassau grouper the origin of the dorsal fin, and in the nostri1 s are subequal. The Nassau having 12, sometimes 13, soft anal-fin grouper can be distinguished from rays (8-9 in Epinephelus spp.). It Mycteroperca spp. by its less elon- differs from other species of gated body and eight soft anal fin Mycteroperca in having a gently round- rays. Other serranid species have ed preopercle, with no definite lobe less robust bodies, soft dorsal and anal fin bases that are not covered by fishes. The body generally is scales or thick skin, and 10 or fewer uniformly brownish red with a lighter dorsal fin spines. ventral coloration and a transient pattern of whitish spots. While inac- tive it may have a banded pattern Color and Pigmentation that matches its surroundings and is similar to that of the Nassau grouper. As a rule, groupers can undergo rapid and dramatic changes in color and pattern, which are under nervous Nassau grouper. In this species the and hormonal control. The extent of body qround color can be tawn.~to change depends on their surroundings pinkish and red, with five dark verti- and the grouper's own activity. Life cal bars; the third and fourth bars color can be used to distinguish dif- divide above the lateral line and ferent species, but specific color and their adjacent branches join to form a distinctive markings such as bars and w-shaped mark. The Nassau and red spots fade soon after the fish dies, groupers are somewhat similar in gene- and undergo further discoloration when ral appearance, but can be distin- the fish is preserved in Formalin or guished because the Nassau grouper has alcohol. a black saddle on top of the caudal peduncle, distinctive black spots The following coloration and pat- below and behind the eyes, and a tern descriptions were compiled from characteristic "tuning fork" shaped Rivas (1964), Cervigon (1966), Bohlke mark on top of the head, all of which and Chap1 in (1968), Fi scher (1978), are lacking in the red grouper. and the Source Document for the Snapper-Grouper Fishery of the South Moe (1963) reported the col lec- Atlantic Region (1983). tion of a xanthic specimen of red grouper off Fort Myers, Florida, that Black grouper. 'This species can agreed in general morphology and change its body coloration and color meristic counts with published pattern over a wide range, from light descriptions of the red grouper, and tan or gray ground color with rows of appeared normal in all respects except rectangular darker blotches, to dark coloration. Overall body coloration reddish gray with short dark bands, to was a brilliant orange-yellow on the whitish with a few dusky bars on the dorsal and upper lateral surfaces, fin margins. In general appearance the shading into reddish-pink on the lower black grouper resembles the yellowfin lateral and ventral surfaces. grouper (M. venenosa) and the gag (M. microlepis], but the black grouper can be distinguished by its straighter posterior caudal margin and rows of Black groupers can reach total rectangular dark blotches on the body. lengths greater than 1 m and weights Blotches are larger and more over 65 kg, but most of those caught quadrangular and regularly aligned in range up to 70 cm TL and about 26 kg the black grouper than in the yellow- in weight. During the 1977 and 1978 fin grouper; in addition, each pecto- Metropolitan South Florida Fishing ral fin of the black grouper has a Tournaments, several black groupers of narrow orange margin, whereas each of 22 to 36 kg were caught in the Florida those in the yellowfin grouper has a Keys (Official Entry Forms, unpubl.). broad yellow margin that sharply con- Male red groupers are reported trasts with the spotted basal portion. up to 72 cm TL and females up to 70 cm TL; most fish in commercial catches Red grouper. This grouper has one range from 45 to 70 cm TL. Nassau of most variable color patterns among groupers can grow to about 1.2 m TL and 20 kg, but most of those marketed nearshore reefs (Beaumariage and weigh 2 to 10 kg (BOhlke and Chaplin Bullock 1976). It has been reported 1968; Fischer 1978). to depths of 189 m, at bottom tempera- tures from 15 to 30 OC, but most are collected at 19 to 25 OC (Roe 1976). LIFE HISTORY The Nassau grouper is generally Habitat found near high-relief coral reefs and rocky bottoms. from the shoreline to a Like most species of groupers, depth of at least 90 m (Fischer 1978; the black, red, and Nassau groupers Bannerot 1984). Cervigon (1966) re- are secretive, occupying caves, ledges ported that large fish occur at depths and crevices on reefs and shipwrecks greater than about 50 m. In Bermuda, (Smith 1961); the larger fish general- Bardach (1958) reported a marked size ly occupy the lower part of the depth segregation: fish less than 40 cm SL range (Thompson and Munro 1978). were on inshore banks and larger ones were on offshore banks. The species The black grouper lives over may be resistant to changing salini- rocky bottoms, coral reefs, and drop- ties: in the old New York Aquarium, off walls; fish longer than about 65 several Nassau groupers reportedly cm TL are generally restricted to lived for several years in water that water deeper than 20 m and small young occasionally became almost fresh and of the year are usually in shallower was rather pol luted (Townsend 1905). water (Fischer 1978). During monthly collections on a shallow grass flat on Sex Reversal and Spawning Matecumbe Key, Florida, Springer and McErlean (1962) reported collecting Most serranids are protogynous what appeared to be two young black hermaphrodites (fish are first females groupers (20 and 24 rnm in standard and then change into males), and the length, SL) . In Venezue 1a, Cerv igon sex of an individual cannot be (1966) reported that black groupers up accurately determined unless it is to 350 cm SL frequently occur over ripe. There is differential distribu- muddy bottoms of mangrove-fringed tion by sex in black and Nassau grou- coastal lagoons, and that fish longer pers, but not in red groupers than 65 cm SL are generally at depths (Bannerot 1984). greater than about 20 m. Moe (1966) reported this species to a depth of Smith (1959) reported histologi - 151 m, at bottom temperatures of 16 to cal and field observations that 28 OC (mean 20 OC). strongly suggested protogynous her- maphrod it ism in b 1ack groupers. The The red grouper occurs mainly black grouper has been reported to over rocky bottoms, but also lives spawn off Puerto Rico in February over muddy bottoms (Cervigon 1966; (Erdman 1956), and has been observed Fischer 1978). In the eastern Gulf of in spawning condition in the Campeche Mexico it has been reported as occur- Bank area in July and August (Smith ring only over rocky reef bottoms at 1961). Off Bermuda, spawning extends depths of 3 to 122 m, frequently oc- from May to early August; females may cupying crevices, ledges, and caverns weigh up to 22.7 kg, but larger fish in limestone reefs (Moe 1969). Juve- are usually males (Smith 1971). niles may be widely dispersed over hard bottoms at depths of at least 37 Red groupers change from female m, being extremely cryptic and thus to male between the ages of 5 and 10 relatively invulnerable to most col- years, at a rate of about 15% annual- lecting gear. Fish 1 to 6 years old ly. The transition occurs at any (under 50 cm SL) commonly inhabit length greater than about 275 mm SL, but is most common after 500 mm SL is tanks, with no formation of mating reached. Females are mature at 4-6 pairs (Manday and Fernandez 1966). years, but reach their greatest repro- ductive potential at 8-12 years of age. Males do not compose more than Egqs and Larvae 10% of a year class until after age 9 (over 500 mm SL), and the sex ratio is Groupers produce planktonic eggs not equal until about age 15, or 625 that are ferti1 ized external ly, and mm SL (~oe1969; Beaumariage and predators and currents affect their Bu 1lock 1976). Males are reproductive- survival (Smith 1961; Colin 1982). ly significant in the population in Fecundity has bee^ estimated to be age groups 10 years old and older over 5 million eggs at 805 mm SL in (Beaumariage and Bul lock 1976). black groupers; over 785,000 at 445 mm SL for Nassau groupers (Smith 1961); Female red groupers usually do and. 1,469,000 (range 312,000- not spawn until they are about 450 rnm 5,735,700) for red groupers 495 to 667 SL. Off the west coast of Florida, mm SL (Moe 1969). Calculated individ- spawning peaks in April and May in ual egg weights ranged from 12 to 22 waters 20-90 m deep and at water tem- m9. peratures between 19 and 21 OC. Gona- dal activity has been observed as Moe (1969) described the eggs of early as January, and culminates in the red grouper as having no filaments spawning in late spring. The only or other appendages, containing an oil environmental factor that correlates droplet, and beiug less than 1 mm in significantly with gonadal development diameter . Manday and Fernandez ( 1966) is photoperiod (Moe 1969). reported that the eggs of the Nassau grouper were between 0.9 and 1.024 mm in diameter, and described the spe- The Nassau grouper changes from cies' embryological and larval devel- female to male at a length between 300 opment until the yolk sac was and 800 mm, which presumably indicates absorbed. that an individual fish can spend more than one spawning season as a func- The distribution of grouper lar- tional female. Off Bermuda, 3.2 kg vae is poorly known. Larval red grou- fish are just reaching maturity per probably leave the plankton and (Bardach and Menzel 1957). Spawning become benthic at about 20-25 mm SL near Bermuda occurs between early May (Beaumariage and Bullock 1976). and mid-August (Smith 1971). In the Caribbean, ripe fish were collected Johnson and Keener (1984) re- between February and May, and sexually ported that the different patterns of inactive ones were collected in serration on the long second dorsal October and November (Munro et al. and pelvic fin spines that charac- 1973). Nassau groupers form spawning terize grouper larvae are consistent aggregations of as many as 100,000 features that can be used to identify fish off Bimini, Bahamas, for 1 week larvae of certain species groups. during the full moon in January (Smith These patterns can identify larvae of 1972). Similar aggregations have been some grouper species at lengths as reported by Miller (1984) in ; short as 5-6 mm, in lieu of ranges of by Bannerot (1984) off the southern meristic counts, where considerable Berry Islands, Bahamas (both also overlap exists among many American during the full moon in January) ; by grouper species. In the genus Burnett-Herkes (1975) off Bermuda; and Mycteroperca, which includes the black by Olsen and LaPlace (1978) in the grouper, species separation of the Virgin Islands. Nassau groupers have larvae based on spinelet morphology is been observed to spawn at night in not possible. However, larvae of the genus can be distinguished from larvae common in seagrass beds (Randall of other grouper genera by the fol- 1968). lowing characteristics: "higher number of anal soft rays (10-13, usually 11, Movement and Migration vs. 7-10, usually 8 or 9), character- istic spine morphology.. .and presence Most grouper species appdrent ly of a cleithral pigment spot." Larvae migrate vertically as they grow, the of the red group.er, the red hind larger fish living at progressively (Epinephelus guttatus), and the greater depths. However, adult grou- speckled hind (E. drummondhayi) can be pers may stay in the vicinity of spe- separated from larvae of other cific reefs for long periods. Black epinepheline species with the excep- groupers tagged and released in in- tion of the mutton hamlet (E.. afer), shore reefs in the Florida Keys dis- but cannot be distinguished among played strong home-reef specificity, themselves because they have simi lar and "even a hurricane failed to dis- body form, spinelet morphology and rupt their residence" (Beaumariage and relative spine length. Larvae of the Bullock 1976). Nassau grouper and the rock hind (E. adscensionis) can be separated from Using evidence from tag and re- all other western Atlantic grouper capture studies of red groupers, Moe larvae by their fin ray counts, with (1969) summarized their offshore move- the exception of the jewfish (E. ment. During their early years they itajara), which has a particular spine remain in shallow water, usually 3 to morphology. 18 m deep. Later, at about 400-450 mm SL and 4-6 years, they leave the near- The function of the spinelets is shore reef environment and move to not known. According to Johnson and depths greater than 36 m. Evidence Keener (1984) they may be involved in supporting the tagging and recovery interspecific recognition, but this is data was given by size and age not 1ikely because the spinelets are analyses of onshore and offshore relatively transparent and lack pig- fishes. The offshore movement seems to mentation. The purpose of these struc- coincide with the onset of sexual tures may be to repel or hook maturity. Commercial fishermen have attacking predators. reported seasonal movement of adults in offshore waters 27 to 91 m deep; some school ing or group movement among Juveni les adults has been evidenced by commer- cial catches of tagged individuals Juveniles of the three species of (Source Document for the Snapper- groupers treated here are commonly Grouper Fishery of the South Atlantic found in inshore seagrass beds in Region 1983). A1 though no alongshore south Florida, and are often caught in migrational patterns were noted, a few shal lower waters than those occupied individual red groupers moved 28 to 72 by adults. Juvenile red groupers do km from tagging locations. not wander far away from their reefs of residence and are scattered in low Beaumariage and Bullock (1976) densities over hard bottoms in water reported that tagged Nassau groupers at least 36 m deep, where they are in the Florida Keys, transported to difficult to collect due to their other nearby reefs, promptly returned cryptic behavior (Moe 1969). Juveni le to their original reef, showing a red groupers are "exact miniatures" of "strong home-reef specificity. " the adults in form and color, and they Randal 1 (1962) reported that tagged were often collected in seagrass beds Nassau groupers in the Virgin Islands in water shallower than 15 m (Smith moved at most about 820 m from the '1971). Small Nassau groupers are also site of release. From results of tagging studies, Springer and McErlean FISHERIES (1962) reported that the Nassau grouper tends to become established in Recreational and Commerci a1 Fisheries an area. Estimated grouper 1andings and Nassau groupers have been report- fishing effort from 1983 to 1986 by ed to form enormous spawning aggrega- U.S. sport fishermen are sown in Table tions and undergo spawning migrations. 2. Recreational grouper fishermen use A large aggregation of Nassau groupers a variety of boats in pursuit of their was observed in the Bahamas by Smith sport. Boats are a necessity since (1972). The fish congregated as a grouper catches from shore and dock school of perhaps as many as 100,000 fishing are rare. Boats used vary from in water about 27 m deep. Analyses of 3.7 m long, privately-owned skiffs with gonad samples indicated that fish of outboard motors to "party boats," also both sexes were present and ripe. Some known as head boats, up to 26 m long individuals in the aggregation dis- and powered by diesel engines. Party played an unusual color pattern; they boats take groups of fishermen to were dark above and light below and fishing reefs for a fee. Boat numbers their characteristic head markings have increased greatly recently. were reversed. Simi lar spawning aggre- Conventional types of fishing rods and gations were observed by Bannerot reels are used routinely, and occasion- (1984), Mi1 ler (1984), and others. ally electric reels are used. Minor catches are made by spear fishermen, who most often use scuba gear. Fish traps were used sporadically in Florida beginning in 1919 (Schroeder 1924). GROWTH CHARACTERISTICS They were prohibited in Florida State waters in 1980, but are still legal to Mean back-calcul ated total lengths use in a1 1 Federal waters. for black groupers aged 1, 5, 10, and 14 years were 260, 664, 975, and 1,110 Since 1948, commerci a1 grouper mm (Manooch and Mason 1987). Growth in 1andings have been significantly 1arger length was most rapid for the first 3 in Florida than in several other States to 4 years, and then gradually slowed. combined (Figure 3). Allen and Tashiro (1976) estimated that about 96% of The rate of growth of male and commercial grouper catches were made female red groupers is be1 ieved to be with hand1 ines. Incidental catches similar, though males reach a larger were made in shrimp trawls, spiny ultimate size than females. Their lobster traps, fish pots, haul seines, effective fishable life span is about trammel and gill nets, and longlines. 17 years, but life span may reach 30 Long1 ines became a major gear ( in terms years (Moe 1969; Beaumariage and of pounds landed) in the gulf grouper Bullock 1976). Data from the Schlitz fishery in the early and mid-1980's and tagging program indicated that the red accounted for the increase in grouper grows at about 5.8 to 10.3 mm commerci a1 1andings then (Goodyear per month (Moe 1966, 1967). For Nassau 1988). Vessels are usually 7.9 to 24 m groupers in the Virgin Islands, and long, and frequently use a small based on results from tagging studies, steadying sail while fishing over Randall (1962) reported the following reefs. Some boats are mu1 ti-purpose, mean growth rates: fish 175-250 mm TL used in shrimp and lobster fisheries grow about 4.55 mm/month; 251-325 mm TL when not used for groupers. about 3.5 mm/month; and 326-451 mm TL about 1.92 mm/month. Table 1 presents Additional information on the reef theoretical growth parameters for red fish fisheries in the gulf is provided and Nassau groupers. by Waters (unpubl .) . Fishermen, 1i ke Table 1. Yield-per-recrui t parameters for black, red, and Nassau groupers. N.D. = no data available; t = age at length of first capture; to= age at length 0; K = von Bertalanffy coefficient; Z = total mortality; Loo = length at infinity; TL = total length; SL = standard length.

------Length- weight Species t, to K Z Loo relationship Area Source

Black 5-7 -0.927 0.116 0.49-053 1352 ~=5.55x10-~TL~.'~' Mostly Manooch and South Mason (1987) Florida Red 1 -0.449 0.179 0.322 672 ~=4.34x10-~SL~.~~ West Moe (1x9) Florida Nassau 4 -0.488 0.185 N.D. 974 W=0.1393 SL~." U.S. Olsen & Virgin LaPl ace Is1ands (1978)

the fishing vessels they use, may work The strong import market for groupers, only part time fishing for grouper, which are imported into south Florida often working in other fisheries or from throughout the Caribbean and from jobs. as far away as Brazil, implies a weak export market. Manooch and Mason (1987) present a yield-per-recrui t fishery model for Reuul ations black grouper. The minimum size for any species Marketinq (including species such as scamp and gag) is 18 inches total length. New Data from a telephone survey gear restrictions prohibit use of reported by Cato and Prochaska (1976) longline nets and stab nets (also showed that Gulf of Mexico groupers called sink nets) off the Atlantic were usually sold by fish dealers fresh coast of Florida. and iced (82% by volume); 16% were filleted and only about 2% were frozen whole. The principal markets are in Artificial Reefs the Southeastern United States, relatively close to the important Artificial reef programs deslgned fishing grounds, thereby permitting to improve recreational fishing in the shipments of a fresh, iced product. ocean have become popular in recent Most dealers sold their fish to other years; most activity in United States wholesalers (52%); about 35% went to waters has been in Florida. Artificial retail markets or agents. The rest reefs consisting of a wide variety of were sold to New York agents. materia1 s, ranging from abandoned vessels to designed and prefabricated Groupers probably do not contribute structures and even junk (debris, 1arge substantially to the export market; appl iances, etc. ) , have been placed on perhaps only 1% by volume of both the coasts of Florida (greatest groupers and snappers are exported. activity has been in Dade County). Table 2. Recreational statistics (catch and effort) for groupers in Florida, 1983- 86.'

Florida (east coast) Florida (west coast) Number of Effort (nymber Number of Effort (nymber Year fish caught of trips) fish caught of trips)

- - - - '~romNational Marine Fisheries Service (1984-87). 2~ripsfor all sport fish.

A controversy exists as to the best variety of fishes and crustaceans depths for artificial reefs. Among the (Thompson and Munro 1978) during the recreational fishermen, there seems to day and at night. Generally, feeding be a consensus favoring water about is most active at dawn and dusk 73 m deep, where diving and spear- (Randal 1 1967). Foods include fishing are discouraged and reefs tend cephal opods, crustaceans and other to attract large fish. The long-term invertebrates, and fishes. By opening effect of artifici a1 reef placement the mouth and rapidly dilating the gill would seem to be that recreational covers to draw in water, groupers can catches of groupers would improve, generally engulf prey who1 e (Bardach et perhaps in some proportion to the a1 . 1958). The usual collection method numbers and kinds of artificial reefs (baited hand1 ines) may bias results of instal led (Cardozo and Hirsch 1985). food studies in groupers because these It is not known if artificial reefs fish frequently regurgitate while being increase fish populations or just brought to the surface and may also redistribute them (Bohnsack, in press). contain fishing bait (Randall 1967; Moe 1969). has been regarded as the collection method most 1ikely to Contaminsnts assure unbiased evaluations of stomach contents (Randal 1 1967). Red grouper from the Southeastern United States had an average of 0.008 ppm DDT and undetectable levels of Black groupers are less intimately PCB's. Levels for black grouper were associated with the bottom, and have 0.009 ppm DDT and up to 0.059 ppm PCB's more slender bodies and better compared to other fish, low levels of developed canine teeth than do the red these contaminants are correlated with or the Nassau groupers, suggesting a low lipid content (Stout 1980). predominately piscivorous diet (Randall 1967). Reported stomach contents ECOLOGICAL ROLE include clupeoid fishes (Cervigon 1966), grunts, and cornet-fi sh (Randal 1 Food and Feedinq Habits 1967). Both black and red groupers have been reported to prey on pink Groupers are unspecialized and shrimp, Penaeus duorarum (Cos tello and opportunistic carnivores, feeding on a A1 len 1970). Florida. Alabama. Other states.

1949 1954 1959 1964 1969 1974 1979 1984 Year

Figure 3. U.S. grouper landings (primarily dressed weights) by States. "Other States" includes North Carolina, South Carol ina, Georgia, A1 abama, Mississippi ,Louisiana, and Texas. From A1 len and Tashiro (1976).

Off the Dry Tortugas, Florida, red their better catches during day1 ight. groupers were reported to feed on Cuban commercial boats fishing in the lbtjanid and sparid fishes (Gudger Campeche Bank area reported catching 1929) and on various other fishes, more and larger red groupers at night octopuses, shrimps, stomatopods, and than during the day (Zupanovich and spiny lobsters (Longl ey and Hildebrand Gonzalez 1975). 1941). Randal 1 (1967) reported unidentified crustaceans, , and Randall (1965) reported that the fishes in the stomachs of two specimens stomach contents of 150 Nassau groupers collected off the Virgin Islands. In collected during daylight off the the stomachs of red groupers collected Virgin Islands and Puerto Rico by commercial and sport fishermen off consisted of about 53% fish, 39% the Florida west coast, Moe (1969) crustaceans, 5% cephalopods, 2% found small fish of several species, gastropods, and 2% pelecypods. The octopuses and squids, crabs most abundant families of fish in the (particularly msspp. and Cal a~~adiet, 1isted in order of occurrence, spp.), panulirid and scyllarid were parrot-fi shes (Scaridae), wrasses lobsters, shrimps, and unidentified (Labridae), damsel fishes crustaceans. Moe (1969) also indicated (Pomacentridae), squirrelfishes that red groupers seemed to feed more (Hol ocentridae) , snappers (Lutjani dae) , on invertebrates, particularly and grunts (Haemul idae) . Crustaceans crustaceans, than on fish, but that dominated the diet of smaller fish, feeding habits may change in large whereas fish predominated in the groupers, which probably consume a stomach contents of 1 arger Nassau greater proportion of f ish. Longl ey groupers (over 300 mm SL). Crustaceans and Hildebrand (1941) reported no included crabs (Majidae, Portunidae, difference in feeding habits of red Cal appidae, Porcell anidae, and groupers between day or night, but Moe Xanthidae), stomatopods (Squill idae) , (1969)indicated that fishermen on the hermit crabs (Paguridae), panul irid west coast of Florida generally made lobsters, and caridean and penaeid shrimp. Moll usks incl uded octopuses and preyed upon by sharks and other large squids, arks (Arcoida), and queen fishes. Large groupers are probably conchs (Strombus giqas) , . Randal 1 preyed upon only by certain shark (1964) suggested that despite their species. Only two species of sharks lacking the dentition to cru!!h the were reported by Compagno (1984) to she1 1s, Nassau groupers may consume prey on groupers: the sandbar shark queen conchs after other predators have (Carcharhinus ~lumbeus) and the great made soft parts avail able. In Bahamian hammerhead (S~hvrna mokarran) , but waters, however, Nassau groupers have other species of sharks undoubtedly been observed forcefully pull ing conchs feed on groupers. out of their shells (B. Rose, commerci a1 fisheman, Freeport, Bahamas; Parasites and Diseases pers. comm. ) Moray eels (Muraenidae) have been reported in stomachs of Black, red, and Nassau groupers are Nassau groupers collected in the Virgin hosts for a number of parasites (Table Islands and Puerto Rico (Randall 1965) 3). The effect of these parasites on and Jamaica (Thompson and Munro 1978). the health of their hosts is unknown. Cervigon (1966) reported that Nassau Groupers from the Tortugas Is1ands, groupers in Venezuela fed mostly on Florida, were hosts for several species crustaceans. of digenetic trematodes (Manter 1947). Overstreet (1968), who surveyed species Comuetitors and Predators of fish in Biscayne Bay, Florida,repor- ted digenetic trematodes from black Interspecific competition for food and Nassau groupers. In waters off and shelter among groupers is likely Campeche, Mexico, Fajer et al. (1979) because of the overlap in habitat, found larval cestodes and nematodes in distribution, size, and food habits red groupers, and noted that males (Thompson and Munro 1978). Adults of tended to have more parasites than most grouper species occupy high 1eve1 s females. They also observed that the in the complex trophic webs of reef 1arger groupers (specimens that ranged communi tie's. These webs include many from 320 to 500 mm TL) tended to be species of general ized, opportunistic more heavily parasitized than small er predators (Bohnsack 1982; Bannerot ones--an observation that has been made 1984), including groupers, which can be for a wide range of host species. assumed to constantly compete with, and prey upon, similar species. Other Observations on grouper parasites competitors for food probably include were made by Thompson and Munro (1978) various species of jacks (Carangidae) , from specimens caught around Jamaica. snappers ( Lutjanidae) , barracudas Parasites were not identified and only (Sphyraenidae), and sharks general groups were reported. In (Carcharhinidae, Sphyrinidae). Nassau groupers, they found parasitic isopods in the nostrils, encysted According to Thompson and Munro larval tapeworms commonly in the (1978) "nothing is known of the causes viscera, and nematodes in the ovaries. of natural mortal ity in groupers. " They observed, "Heavy infection by this Small, cryptic individuals probably (nematode) parasite can drastical ly fall prey to fishes such as moray eels reduce the number of eggs produced by (Muraenidae), which can attack them in an individual ." The gonads of male their hiding places. Large groupers of fish were not infected. several species are also reported to readily feed on smaller groupers (Smith The only report on grouper diseases 1961). Thompson and Munro (1978) that we found in the literature was an mentioned the 1ack of pub1 ished account by Moe (1969) of a large information on grouper predators, and skeletal tumor removed from a red suggested that groupers are probably grouper. The fish was caught about Table 3. Parasite species in black, red, and Nassau groupers.

Species of Location Geographic parasite in host location Source

BLACK GROUPER

Digenetic trematode

Lecithochirium microstomum Stomach Biscayne Bay, F1 Overstreet (1968) 11 II 11 Intestine 11 I1 Intestine & I1 I1 pyloric caeca

RED GROUPER

Digenet ic trematodes

He1 icometra torta N.D. Tortugas, F1 Manter (1947) Lepidapedon levenseni Intestine & I1 11 caecum Stephanostomum dentatum Intestine I1 I1

Cestoda

Callotetrarhynchus sp. Muscles Campeche Bank Fajer et al. ( larvae-encysted) Digestive tract (Mexico) (1979) Liver Abdominal cavity Gonads

Nematoda

Anasak is Digestive tract ( larvae-encysted) Liver Gonads

NASSAU GROUPER

Digenet ic trematode

Helicometra torta N.D. Tortugas, F1 Manter (1947) I1 Pyloric caeca Biscayne Bay, F1 Overstreet (1968) Lecithochirium microstomum Stomach Biscayne Bay, F1 Overstreet (1968) Leci thochirium parvum II Tortugas, F1 Manter (1947) Sterrhurus muscu lus at Biscayne Bay, F1 Overstreet (1968) a~.~.Smith (1961) reports 0 istho orus epinephely occurring in red grouper, based on a report by Manter *record of this trematode was found in Manter's report. 43 km southwest of Venice, Florida, in Behavior May 1967. No information on length, age, or sex of the fish was recorded. Groupers can generate sound in Otol ith abnormal growths have been different ways, particularly by reported in red groupers by Moe (1969) vibrating their thin-walled swim and in Nassau groupers by Thompson and bladder by sudden contraction of axial Munro (1978). muscles, such as the single pair of bi1 ateral muscles behind the opercl es. Ciquatera Various sounds have been attributed to all three species: low-frequency, Ciguatera is a toxin transmitted by high-amp1 itude "mooing-1 iken sounds, hundreds of species of tropical fishes. deep booms, and sustained rumbles, It causes gastrointestinal, vibrant grunts (single- or in rapid cardiovascul ar, and neurological series), and long noises from grinding disturbances in persons who eat toxic of teeth when fighting for food. Sound fish, resulting in prolonged disabi 1i ty may have many functions, including and long and expensive recovery warning, intimidation, orientation, and periods. Several species of Western recognition (Hazlett and Winn 1962; At1 antic groupers, including the back Fish and Mowbray 1970). Nassau and the red groupers, are routinely groupers are diurnal or crepuscular in reported to be ciguatoxic, whereas their movements (Col 1ette and Ta'l bot others such as the Nassau grouper seem 1972) and do not usually move far from to be uniformly nontoxic, throughout cover (Starck and Davis 1966). They their range. In south Florida are often wary of divers and quickly ciguatera is endemic, and of all retreat into hiding places, producing grouper species the black grouper is a series of grunting sounds (Hazlett the most often implicated; this species and Winn 1962). Black and Nassau is considered toxic throughout the groupers were reportedly attracted by Caribbean, except near St. Croix, U.S. irregularly pulsed signals during Virgin Islands (de Sylva and Higman experiments designed to attract and 1980; Pol i 1982). film sharks off Bimini, Bahamas. The groupers approached slowly, came to The occurrence of the toxin is rest in front of the sound projector, usually restricted to 1arge predatory and slowly moved away after 10-30 species associated with reefs. The seconds (Myrberg et a1 . 1969). Randal 1 larger older fish are generally more (1962) reported that Nassau groupers toxic than small younger fish because have a tendency to re-enter fish traps; the old fish have had time to some tagged fish were recaptured accumulate the toxin. Even within several times, presumably re-entering certain species the distribution of to feed on other fish in the traps. toxic fish is not geographically continuous, occurring in some areas and Nassau groupers are often involved not in others (Olsen et al. 1984). In in symbiotic cleaning behavior at a study based on interviews (Poli "cleaning 'stations" on coral reefs. 1982), "groupers" were responsible for Species most often reported involved the largest number of recorded cases of include several species of gobies ciguatera intoxication in Florida (Gobiosoma spp.) and shrimp that mainly during two periods, "pre-1978" and remove gnathiid isopods from the 1978-1980. Many times the species bodies, fins, gill chambers and mouths responsible were not identified due to of groupers (Bohlke and McCosker 1973; the processing procedures used for Darcy et al. 1974; Sargent and marketing the fish. Wagenbach 1975).

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Hazlett, B., and H.E. Winn. 1962. Moe, M.A., Jr. 1966. Tagging fishes Sound producing mechanism of Nassau in Florida offshore waters. Fla. grouper, E ine helus striatus Bloch Board Conserv. Tech. Ser. No. 49. 40 Copeia 196- PP Johnson, G.D., and P. Keener. 1984. Moe, M.A., Jr. 1967. Prolonged sur- Aid to identification of grouper vival and migration of three tagged larvae. Bull. Mar. Sci. 34(1):106- reef fishes in the Gulf of Mexico. 134. Trans. Am. Fish. Soc. 96:228- 229. Longley, W.H., and S.F. Hildebrand. 1941. Systematic catalogue of the Moe, M.A., Jr. 1969. Biology of the fishes of Tortugas, Florida, with red grouper (Epinephelus morio observations on colour, habits and Valenciennes) from the eastern local distributions. Carnegie Inst. Gulf of Mexico. Prof. Pap. Ser. Wash. Pap. Tortugas Lab. 34. 331 pp. Mar. Lab. Fla. No. 10. 95 pp.

Manday, D.G., and M. J. Fernandez. Munro, J.L., V.C. Gaut, R. Thompson, 1966. Desarrollo embrionario y pri- and P.H. Reeson. 1973. The meros estados larvales de la cherna spawning seasons of Caribbean reef criolla, Epinephelus striatus fishes. J. Fish Biol. 5:69-84. (Bloch) (Perciformes: Serranidae). st. 1nst. Oceanog. Habana ~o.l:35- Myrberg, A.A., Jr., A. Banner, and 45. J.D. Richard. 1969. Shark attrac- tion using a video-acoustic system. Manooch, C.S., 111, and B.L. Mason. Mar. Biol. 2:264-276. 1987. Age and growth of the warsaw grouper and black grouper from the Olsen, D.A., and J.A. LaPlace. 1978. A southeast region of the United study of a Virgin Islands grouper States. Northeast Gulf Sci . 9:65-75. fishery based on a breeding aggrega- tion. Proc. Gulf Caribb. Fish. Inst. Manter, H.W. 1947. The digenetic 31 :130-144. trematodes of marine fishes of Tortugas, Florida. Am. Midl. Nat. Olsen, D.A., D.W. Nellis, andR.S. 38(2) : 257-416. Woods. 1984. Ciguatera in the eastern Caribbean . NOAA/NMFS Mar. May, R.M., J.R. Beddington, C.W. Fish. Rev. 46(1) :13-16. Clark, S.J. Holt, and R.M. Laws. 1979. Management of mu 1tispecies Overstreet, R.M. 1968. Digenetic fisheries. Science 205:267-277. trematodes of marine fishes from Biscayne Bay, Florida. Unpubl. Miller, W. [1984]. Spawning aggrega- Ph.D. Dissertation. University of tions of the Nassau grouper, Miami, Coral Gables, Fla. 188 pp.

Epinephelus striatus, and ' the associated fishery in Belize. presented at the meeting Advances Poli, M.A. 1982. A review of ciguatera in Reef Science, October 26-28, (tropical fish poisoning), with spe- 1984, University of Miami, Miami, cial reference to the Caribbean and Fla. (unpubl. ms.). an investigation into its incidence and significance in Florida. Unpubl. Moe, M.A., Jr. 1963. Partial albinism M.S. Thesis. University of Miami, in a xanthic specimen of Epinephelus Coral Gables, F1. 56 pp. Randall, J.E. 1962. Tagging reef fi- Smith, C.L. 1961. Synopsis of biolo- shes in the Virgin Islands. Proc. gical data on groupers (Epinephe- Gulf Caribb. Fish. Inst. 14:201-241. lus and allied genera) of the %tern North Atlantic. FA0 Fish. Randall, J.E. 1964. Contributions to Biol. Synop. No. 23. 61 pp. the bioloav of the aueen conch. Strombus 7"as Bull. ~ar.Sci. ~ulf Smith, C.L. 1971. A revision of the Cari bb. lw~46-295. American groupers: Epinephelus and allied genera. Bull. Am. Mus. Nat. Randall, J.E. 1965. Food habits of the Hi st. 146 :67-242. Nassau grouper (Epinephelus stria- &). Assoc. Isl. Mar. Labs. Caribb., Sixth Meeting:13-16. Smith. C. L. 1972. A s~awninqaqqre- on of Nassau grouper, Epi ieph- Randall, J.E. 1967. Food habits of elus striatus (Bloch). Trans. reef fishes of the West Indies. Am. Fish. Soc. 101: 257-261. Stud. Trop. Oceanogr. Univ. Miami 5 :665-847. Smith, G.B., H.M. Austin, S.A. Randa 11, J. E. 1968. Caribbean reef Bortone, R.W. Hastings, and L.H. fishes. TFH Publ ications, Jersey Ogren. 1975. Fishes of the Florida City, New Jersey. 318 pp. Middle Ground with comments on ecology and zoogeography. Fla. Mar. Rivas, L.R. 1964. Western Atlantic Res. Publ. No. 9. 14 pp. serranid fishes (groupers) of the genus E ine helus. -Q. J. Fla. Acad. sci. 2- Source Document for the Snapper- Grouper Fishery of the South Roe, R.B. 1976. Distributions of Atlantic Region. 1983. South snappers and groupers in the Gulf Atlantic Fishery Management Council, of Mexico as determined from Charleston, South Carolina. 306 exploratory fishing data. Pages PP. 129-164 fi H. R. Bullis and A.C. Jones, eds. Proceedings: colloquium on snapper-grouper Springer, U.G., and A.J. McErlean. fishery resources of the western 1962. Seasonality of fishes on a central Atlantic Ocean. Fla. Sea south Florida shore. Bull. Mar. Grant Rep. 17. Sci. 12(1) :39-60.

Sargent, R.C., and G.E. Wagenbach. 1975. Cleaning behavior of the Starck, W.A., 11, and W.P. Davis. shrimp Periclimenes antho hilus 1966. Night habits of fishes of (Crustacea: Decapoda, -+Natantia Alligator Reef, Florida. Icthyol . Bull. Mar. Sci. 25(4):466-472. Aquarium J. 38(4):313-356. Schroeder, W.C. 1924. Fisheries of Key West and the clam industry of Stout, V.F. 1980. Organo-chlorine southern Florida. Rep. U.S. Comm. residues in fishes from the northeast Fish. for 1923, App. 12, U.S. Bur. Atlantic Ocean and Gulf of Mexico. Fish., Doc. No. 962. U.S. Natl. Mar. Fish. Serv. Fish. Bull. 78:51-58. Smith, C. L. 1959. Hermaphroditism in some serranid fishes from Bermuda. Pap. Mich. Acad. Sci., Arts, Lett. Thompson, R., and J.L. Munro. 1978. 44:lll-118. Aspects of the biology and eco- logy of Caribbean reef fishes: Waters, J.Y. [1988.] Review of the Serranidae (hinds and groupers). reef fish fisheries in the Gulf of J. Fish Biol. 12:115-146. Mexico. 89 pp. Unpubl . MS.

Zupanovich, S., and P. Gonzalez. 1975. Townsend, C.H. 1905. Report to the Investigaciones y pesqueria de la director of the aquarium. Rep. cherna en el Banco de Campeche. N.Y. Zool. Soc. 9(1904):88-103. Mar Pesca 112:22-27. 50171 -101 REPORT DOCUMENTATION 1. REPORT NO. 2. 3- Rec~pient'sAccession NO. PAGE ~iological&port 82(11.110)*~ I I 4. Title and Subtitle 5. Repon Date Species Profiles: Life Histories and Environmental Requirements August 1989 of Coastal Fishes and Invertebrates (South Florida)--Black, Red, and Nassau Groupers

7. Author(s) 8. Perforrnvng Organization Rept. NO. Darryl E. Jory and Edwin S. Iversen I 9. Perterming Organization Name and Address 10. Proiect/Task/Werk Unrt No. 11. Contract(C) or Grant(G) No. I It. Sponrorlng Organization Name and Address U. S. Department of the Interior U. S. Army Corps of Engineers Fish and Wildlife Service Waterways Experiment Station National Wetlands Research Center P.O. Box 631 Washington, DC 20240 Vicksburg, MS 39180

-- 15. Supplementary Notes *U.S. Army Corps of Engineers Report No. TR EL-82-4 I 16. Abstract (Limit: 203 words) Black, red, and Nassau groupers (Mycteroperca bonaci, Epinephelus morio, and E. striatus, respectively) are widely distributed on rocky bottoms and reefs along the south Florida coast. They are the most valuable marine finfish group in Florida, comprising about 25% of the total value of landings in 1984. he three species can be distinguished by morphometric, meristic, and body color characteristics. Younger fish are typically found in shallow, inshore grass beds, and larger, older fish are gener- ally restricted to deep waters. The three species are protogynous hermaphrodites. Sexual transition can occur at any length over about 300 mm SL. An offshore movement apparently coincides with the onset of sexual maturity. Spawning aggregations have been observed throughout the year, but occur mostly between late spring and early summer. Fecundity estimates range from about 800,000 to over 5,000,000 eggs per female. Both the eggs and the larvae are planktonic. Their early life history is poorly known. Larvae probably leave the plankton and become benthic at around 20-30 mm SL. Growth rates range from about 2 to 10 mm/month. The three species are unspecialized carnivores, feeding on a variety of fishes, crustaceans, and mollusks. Interspecific competition for food and shelter may be common because of the overlap in distribution, habitat, size, and food habits. For the three species, a number of predators and parasites have been reported. Both the black and red groupers have been imp1 icated in ciguatera poisonings in south Florida. 17. Document Analysis a. Descriptors Marine fishes Life cycles Parasites Fisheries Growth Spawning Feed habits Competition Predators

b. IdentiRenlOpen-Ended Terms Black Grouper (Mycteroperca bonaci) Red Grouper (Epinephelus morio) Nassau Grouper (Epinephelus striatus)

18. Ava~lab#lityStatement 19. Securvty Class (This Report) 21. NO. of Pages Unl imited Unclassified 2 1 20. Securlty Class nhts Page) P. Prlce ! Unclassifed (See ANSI-239.18) OPTIONAL FORM 272 (4-77) (Formerly NTIS-35) Department of Commerce As the Nation's principal conservation agency, the Department of the Interior has responsibility for most of our nationally owned public lands and natural resources. This includes fostering the wisest use of our land and water resources, protecting our fish and wildlife, preserving the environmental and cultural values of our national parks and historical places, and providing for the enjoy- ment of life through outdoor recreation. The Department assesses our energy and mineral resources and works to assure that their development is in the best interests of all our people. The Depart- ment also has a major responsibility for American Indian reservation communities and for people who live in island territories under U.S. administration.

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