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REFEREFCS CO?y D~ Not Remove from the Librow SEDAR25-RD15 I I c rich rind VJildlife ser\~if:e -. -- Blologlcal Repon 82(11-99) Noiional \"Jetlands Research Cents: July 1989 700 Cajun Dome Boulevcrd TR EL-82-4 Lafayette, Louisiana 73506

Species Profiles: Life Histories and Environmental Requirements of Coastal Fishes and Invertebrates (South Atlantic) BLACK SEA

Coastal Ecology Group Fish and Wildlife Service Waterways Experiment Station U.S. Department of the Interior U.S. Army Corps of Engineers SEDAR25-RD15

Biological Report 82(11.99) TR EL-82-4 July 1989

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

BLACK SEA BASS

BY

Linda P. Mercer North Carolina Division of Marine Fisheries Morehead City, NC 28557

Project Officer David Moran National Wetl ands Research Center U.S. Fish and Wildlife Service 1010 Gause Boulevard Slidel 1 , 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 Wetl ands Research Center Washington, DC 20240 SEDAR25-RD15

This series should 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 can be cited as follows:

Mercer, L.P. 1989. Species profiles: life histories and environmental re- quirements of coastal fishes and invertebrates (South Atlantic)--black sea bass. U.S. Fish Wildl. Serv. Biol. Rep. 82(11.99). U.S. Army Corps of Engineers, TR EL-82-4. 16 pp. SEDAR25-RD15

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 biological 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 applicable. 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 Engi neer Waterways Experiment Station Attention: WESER-C Post Office Box 631 Vicksburg, MS 39180 SEDAR25-RD15

CONVERSION TABLE

Metric to U.S. Customary

Multiply To Obtain millimeters (mm) inches centimeters (cm) inches meters (m) feet meters (m) fathoms ki1 ometers (km) statute miles kilometers (km) nautical miles square meters (m2) 10.76 square feet square ki1 ometers ( km2) 0.3861 square miles hectares (ha) 2.471 acres liters (1) gal 1ons cubic meters (m3) cubic feet cubic meters (m3) acre-feet milligrams (mg) ounces grams (g) ounces ki1 ograms (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 25.40 millimeters inches 2.54 centimeters feet (ft) 0.3048 meters fathoms 1.829 meters statute miles (mi) 1.609 ki1 ometers nautical miles (nmi) 1.852 kilometers square feet (ft2) square meters square mi 1es (mi 2, square kilometers acres hectares gallons (gal) 1i ters cubic feet (ft3) cubic meters acre-feet cubic meters ounces (oz) milligrams ounces (oz) grams pounds (lb) ki1 ograms pounds (lb) metric tons short tons (ton) metric tons British thermal units (Btu) kilocalories Fahrenheit degrees (OF) Cel sius degrees SEDAR25-RD15

CONTENTS Page PREFACE ...... CONVERSION FACTORS ...... ACKNOWLEDGMENTS ...... NOMENCLATURE/TAXONOMY/RANGE ...... MORPHOLOGY/IDENTIFICATION AIDS ...... REASON FOR INCLUSION IN THE SERIES ...... LIFE HISTORY ...... Spawning ...... Fecundity and Eggs ...... Larvae ...... Juveniles ...... Adults ...... GROWTH CHARACTERISTICS ...... FISHERY ...... Commercial Harvest ...... Sport Fishing ...... Management ...... ECOLOGICALROLE ...... ENVIRONMENTAL REQUIREMENTS ...... Temperature ...... Salinity ...... Substrate ...... Depth ...... Contaminants ...... LITERATURECITED ...... SEDAR25-RD15

ACKNOWLEDGMENTS

I am grateful to Charles S. Manooch, 111, Gene Huntsman, and R.O. Parker, Jr., of the National Marine Fisheries Service, Beaufort Laboratory, and C. Wayne Waltz of the South Carolina Wildlife and Marine Resources Department for review- ing the manuscript. SEDAR25-RD15

Figure 1. Black sea bass (from Goode 1884).

BLACK SEA BASS

populations of black sea bass may occur along the Atlantic coast, Scientific name ...... Centro ristis separated at Cape Hatteras, North striata (Linnaeus) (Fig* Carol ina (Mercer 1978). Prmcommon name...... Black sea bass (Robins et al. 1980) Other comnon names...... Blackfish, MORPHOLOGY/IDENTI FICATION AIDS black bass, black will (Smith 1907) Class...... Osteichthyes The fol1 owing description was Order ...... given by Miller (1959) : Dorsal X, 11; Fami 1y...... anal 111, 7; total gill rakers plus tubercles usually 22-28 (20-29). lower Geographic range: The black sea bass, limb usually 14 to 18 (14-19); scales striata striata, oc- large, 1ateral 1ine scales 47 (46-49). miccoast from Body robust, the back somewhat ele- Cape cod; Massachusetts, to Cape vated anteriorly; head 1arge; snout Canaveral , Florida (Figure 2), and moderately pointed; mouth large, occasionally as far south as the obl ique. Spinous and soft dorsal fins Florida Keys (Fischer 1978). A sub- continuous, males having the notice- species, C. s. melana, occurs along ably higher fins; caudal fin rounded the eastzrn andrthern coastal in juveniles, trilobed and with areas of the Gulf of Mexico. Two one upper ray extended in adults. 1 SEDAR25-RD15

NORTH CAROLINA

SOUTH CAROLINA

A TlA N TIC OCEAN

MILES 0

KILOMETERS

Figure 2. Distribution of black sea bass along the South Atlantic coast. 2 SEDAR25-RD15

Breeding ma1 es develop an adipose hump usin baited wire crab traps (Rivers on the nape. 19663. Black sea bass comprises the bulk of the North Carolina charter Color in life: smoky gray, dusky boat catch of reef fishes (Manooch brown, or blue-black above, slightly et al. 1981) and is probably second in paler below; centers of scales pale importance to red porgy (Pa rus blue or white, forming 1ongitudinal in the inshore catch stripes along back and sides; sides WAa headboats (sport fishing sometimes mottled or with dark and boats having a passenger capacity of 1ight vertical crossbars. Breeding 30-75 anglers) (Huntsman 1976). males with vivid hues of fluorescent Management regulations for black sea blue and green around eyes and bass caught in the Fishery Conserva- nape; females are lighter and brownish tion Zone (the area in which the or gray-bl ue instead of blue-black United States asserts excl usive (Hardy 1978). Link (1980) described fishery management authority and which four juvenile color phases: an overall extends seaward from 3 to 200 nm) 1ight grayish phase peppered with were established in the Fishery small dark spots; a dark phase with Management Plan for the snapper- dull white spots; a striped phase with fishery of the South Atlantic a single horizontal dark stripe; and a Region (Anonymous 1983). A minimum barred phase with six vertical bars. size limit of 203 mm TL (8 inches) was adopted for black sea bass on the The Atlantic subspecies of black basis of a yield-per-recrui t analysis sea bass differs from Gulf of Mexico that indicated the harvest is less subspecies in the following details: than the maximum possible. gill rakers and tubercles usually 25 to 26, instead of 22-24; pectoral rays 18 or 19, instead of 17 or 18; jaw LIFE HISTORY usually 16% or 17% of standard length, instead of 18%-19%; black at base Spawning of last dorsal spines distinct in juveniles, rather than indistinct to Black sea bass are protogynous almost absent; dark blotches on jaws hermaphrodites. Most individual s usually absent in young. Bortone function first as females, undergo (1977), who compared the osteology- - of sexual succession, and become three. species of Centro ristis, found functional males (Lavenda 1949). no obvious osteol-3-r oqica ifferences The process of sexual succession was between C. 2. stdata and -C. -s. described from histological studies of me1 ana. the gonads (Lavenda 1949; Reinboth 1965; Mercer 1978; Wenner et al. Other species of Centro ristis 1986). Bands of testicular precursor differ in beinq ver.~pa * e to w ite cells are found in various stages of with seven difhse to' distinct dark development in the ovarian wall . vertical bars on side, fins pale, soft Sexual transition begins in the dorsal and caudal with dark to inky posterior region of the ovary and the spots (Mil 1er 1959). proliferation of testicular tissue proceeds forward.

REASON FOR INCLUSION IN THE SERIES Spawning of black sea bass ex- tends from January to June offshore in The black sea bass is a valuable the South Atlantic region, peaking commercial and recreational fish along from March to May (Cupka et al. 1973; the South At1 antic coast. Comnercial Mercer 1978; Link 1980; Wenner et landings increased sharply after 1960 a1 . 1986). Minor spawning occurs in due to the development of a fishery September and October. Spawning SEDAR25-RD15

occurs later in the northern part of younger ages (Mercer 1978; Waltz et the range. The frequency of occurence al. 1979; Link 1980; Wenner et of transitional individuals increases al. 1986). An increased frequency of after both spawning periods, indicat- males occurs between 200 and 240 mn SL ing that sexual succession is primar- and ages IV and V. Individuals ily a post-spawning process (Mercer undergoing sexual succession ranged in 1978; Wenner et al. 1986). length from 118 to 327 mn SL and from ages I to VIII; modes were at 200 mn SL and age 111. Black sea bass mature between age I and IV (Mercer 1978; Link 1980; Wenner et al. 1986). Wenner et al. Fecundity and Eggs (1986) reported that the percentages of females that were mature were 0 at Fecundity relationships for South age 0, 48.4 at age I, 90.3 at age 11, Atlantic black sea bass were deter- 99.1 at age 111, and 100 at all older mined by Cupka et al. (1973), Link ages. The smallest observed mature (1980), and Wenner et al. (1986) and females and males were 90-110 mn and by Mercer (1978) for Middle Atlantic 100-110 mm SL, respectively (Mercer fish (Table 1). Estimates of fecun- 1978; Wenner et al. 1986). Link dity ranged from 17,000 in a fish of (1980) reported that all females of 108 mn SL (age 11) to 1,050,000 in a ages I and I1 collected at depths fish of 438 mm SL of undetermined age <12 m were imnature, whereas some (Wenner et al. 1986). Fecundity was females of ages 1-111 collected at significantly related to length, depths >12 m were mature, and all age weight, and age. IV females were mature. Black sea bass eggs are pelagic, Sex ratios of black sea bass are 0.9-1.0 mm in diameter and hatch differ significantly from 1:l in favor in about 75 h at 16 "C (Wilson 1891; of females at the smaller sizes and Smith 1907). Eggs from artificially

Table 1. Fecundity relationships for South Atlantic black sea bass, where n=sample size, F=fecundity, SL=standard length (mn) , TL=total length (mm), W= weight (g) , and A=age.

Size Reference n Fecundity relationship r range No. of eggs (mn SL)

Cupka et al. 18 log F = 0.308 + 1.973 log SL 0.67 139-250 29,770-121,500 f.-- 19731 -, log F = 3.349 + 0.629 log W 0.69 log F = 4.543 + 0.784 log A 0.70

Link 18 log F = -2.10 + 3.03 log SL 0.94 137-335 26,938-370,443 (1980)

Wenner et al. 115 log F = -0.309 + 2.318 log SL 0.65 108-438 17,000-1,050,000 (1986) log F = -0.605 + 2.335 log TL 0.62 log F = 3.057 + 0.822 log W 0.65 log F = 4.529 + 0.913 log A 0.33 SEDAR25-RD15

spawned gulf black sea bass (C. s. berg (1972) col 1ected young-of-the- melana) hatch in about 38 h aft& year black sea bass throughout the fertilization at 23 "C under an 11 h the year in the lower reaches of a photoperiod (Hoff 1970). Georgia estuary, mostly over she1 1 bottom. In the St. Johns River, Florida, Tagatz (1967) collected ju- Larvae veniles (28-71 mm FL) in June and July at 14.5 to 24.7 ppt and 26.6 to Little is known of the early life 27.4 "C. He collected larger individ- history of black sea bass. Larvae uals (91-176 mm FL) in February, (2-13 mm) have been collected from March, and November at salinities of June to November between Sandy Hook, 7.7-13.1 ppt and 11.0-17.4 "C. New Jersey, and Cape Lookout, North Carolina (Kendall 1972). The larvae were collected in tows from the Adults surface to a depth of 33 m, 4 to 82 km from shore where water depths Adult black sea bass are found in ranged from 15 to 51 m. The pelagic offshore areas in depths of 10 to 120 existence of this fish is short. m; however, most are at 20 to 60 m Larvae longer than 13 mm SL were not (Struhsaker 1969; Bearden and McKenzie col 1ected, presumably because they 1971; Cupka et al. 1973; Grimes et al. become demersal or estuarine near that 1982; Chester et al. 1984; Sedberry size. It is not known what part of and Van Dolah 1984; Parker and Ross the population remains at sea, nor 1986). are the routes and mechanisms of 1arval transport known. Kendal 1 Black sea bass are migratory in (1972) described armature, body shape, the northern part of their range, development of meristic characters, whereas south of Cape Hatteras, North and pigment patterns of larvae 2.1 to Carolina, they are apparently resi- 13.0 mm SL. dents in given areas year round. In the Middle Atlantic Bight black sea Juveniles bass move inshore and northward in spring and offshore and south in fall, Juvenile black sea bass occur in probably due to changes in temperature saline areas of estuaries along the (Pearson 1932; Nesbit and Neville coast from Florida to Massachusetts 1935; Musick and Mercer 1977). Off (Kendall 1977) and in offshore areas South Carolina the lowest bottom (Cupka et al. 1973). Link (1980) temperatures recorded in the depth reported that young-of-the-year (30-50 range inhabited by adult sea bass are mm SL) appeared on inshore jetties in 10 "C or higher during all seasons late May and early June. Within (Walford and Wicklund 1968). Tagging estuaries, juveniles are found around studies off South Carolina and along jetties, piers, wrecks, and shell the east coast of Florida have bottom such as oyster reefs. indicated that black sea bass there are essentially nonmigratory (Topp Black sea bass have been col- 1963; Beaumariage 1964, 1969; lected year round in North Carolina Beaumariage and Wittich 1966; Moe estuaries at salinities of 1 to 36 ppt 1966; Cupka et al. 1973). A1 though no and temperatures of 6 to 29 "C (Link seasonal movements were observed in 1980; Schwartz et al. 1981). In South South Carolina waters, Cupka et al. Carol ina estuaries juveniles (20-140 (1973) noted that larger specimens rn SL) were found at salinities of were found in deep water, which 8.8 to 37.8 ppt and temperatures of suggests gradual offshore movements 5.6 to 30.4 "C, mainly from July to with increasing age; however, reduced November (Cupka et a1 . 1973). Dahl- fishing pressure offshore away from SEDAR25-RD15

ports may be a factor. Waltz et al. not yet formed; they concluded that (1979) reported that average age the functional ly derived correction increased with depth from 2.4 years at factor used in their study under- 18-26 m to 4.8 years at 37-46 m. estimated the back-cal cul ated 1ength in the younger age groups. Males were larger than females at ages greater GROWTH CHARACTERISTICS than one year.

Age and growth of black sea bass Von Bertalanffy growth parameters have been determined from otoliths are shown in Table 3. The low esti- (Cupka et al. 1973; Mercer 1978; Waltz mates of K (growth coefficient) indi- et al. 1979; Link 1980; Wenner et al. cate that black sea bass attains its 1986). Analysis of marginal incre- maximum size slowly, as do other reef ments indicates that annul us formation fishes (Huntsman and Manooch 1978). occurs in April and May. Ten age Link (1980), who used a logarithmic groups were identified in the South function to describe growth of black Atlantic Bight (Table 2). Mean back- sea bass, obtained values that were calculated lengths at age were similar intermediate to those of other stu- in all studies except that of Cupka et dies. al. (1973) and ages I and I1 of Waltz et al. (1979). The larger size at age Length-weight regressions for found by Cupka et al. (1973) may re- black sea bass were calculated by flect the condition of the population Cupka et al. (1973), Mercer (1978), of black sea bass in the South At- Waltz et al. (1979), and Link (1980) lantic Bight before heavy exploitation (Table 4). No sexual differences in began in 1969. Waltz et a1 .(1979) ob- length-weight re1ationshi ps were found served 37 black sea bass of 45 to 130 in these studies. Slopes and inter- mn SL in which the first annulus had cepts were similar in all studies.

Table 2. Mean back-calculated lengths (mn SL) of black sea bass.

Aoe Reference Sex I I1 I11 IV V VI VII VIII IX X

Cupka et al. Combined 115 164 198 230 265 305 329 (1973)

Mercer (1978) Femal e 93 143 176 199 218 208 Ma1 e 90 145 183 215 238 269 286 294 Combined 87 141 177 205 231 244 Waltz et a1 . Female 74 133 168 190 218 247 257 256 280 330 (1979) Male 73 134 177 210 244 260 277 295 300 340

Link (1980) Femal e 92 142 187 231 247 260 Ma1 e 99 155 202 244 272 277 293

Wenner et al. Combined 88 142 180 212 244 271 283 289 296 303 (1986) SEDAR25-RD15

Table 3. Estimated von Bertalanffy THE FISHERY parameters describing the growth of black seabass. Commerc ia1 Harvest

The fisheries for black sea bass Reference K L , were reviewed by Rivers (1966), Cupka to et al. (1973), Frame and Pearce (1973), and Mercer (1978). In the Mercer -0.183 0.219 352 South Atlantic region black sea bass ( 1978) are mainly harvested by a trap fish- ery; small amounts are caught with Cupka et al. -1.330 0.088 625 trawls and hand 1ines. Smith (1907) (1973) reported that the fishery in the early 1900's was primarily a handline fish- Wenner et al. -0.201 0.231 341 ery. The trap fishery, which devel- ( 1986) oped in 1960, operates year-round along the 18 m contour. Wire-mesh crab pots are used, baited with men- ato= theoretical age at which length haden, herring, squid, mullet, spot, is zero or croaker. Highest landings occur K = rate at which length approaches from early winter to spring due to in- the asymptote creased participation in the fishery Loo = asymptotic length at infinite by shrimp and sport fishermen. Fif- age. teen to 40 traps are set singly over patch reefs. Fishing time ranges from 20 to 45 minutes for each trap. This fishery differs considerably from the trap fishery north of Cape Hatteras, North Carolina, which operates from May to December, and in which 400 to 1,500 unbaited wooden slat traps are set out for the entire season in

Table 4. Weight-length regressions for black sea bass (loglow = a + b [loglO~~] where W = weight [g] and SL = standard length [mml).

2 Reference n a b r

Mercer (1978)

Link (1980)

Cupka et al. ( 1973)

Waltz et al. (1979)

Wenner et al. ( 1986) SEDAR25-RD15

strings of 10 or 20 traps per 1ine, Table 5. Commercial landings (metric with about 200 traps fished daily. tons) of black sea bass in the south Trawl catches of black sea bass are Atlantic region, 1950-1984 (from primarily landed north of Cape Fishery Statistics of the U.S.). Hatteras, North Carolina. Black sea bass are also caught in the handline State fishery for snappers and . Year NC SC GA FL Total Black sea bass landings (Table 5) increased sharply in the South At- 1950 34 115 lantic region after 1960 due to the 1951 43 84 development of the trap fishery. 1952 5 0 45 Cupka et al. (1973) reported that the 1953 3 7 3 5 numbers of vessels fishing for black 1954 19 27 sea bass in South Carol ina increased 1955 9 8 from 8 in 1959 to 62 in 1971. Annual 1956 3 6 15 landings in North Carolina peaked at 1957 16 2 904 t in 1967 and have fluctuated 1958 12 8 since then. South Carolina landings 1959 19 17 were highest in 1970, then declined, 1960 5 7 13 and increased in the early 1980's. 1961 288 147 Georgia and Florida contribute 1 ittle 1962 584 122 to South Atlantic black sea bass 1963 335 120 1andi ngs. 1964 411 106 1965 494 38 Sport Fishi ng 1966 575 62 1967 904 3 0 The recreational catch of black 1968 541 93 sea bass has increased, partly due to 1969 475 327 the development of the headboat fish- 1970 534 351 ery in the mid-1960's (Huntsman 1976). 1971 339 233 The black sea bass is the most abun- 1972 288 248 dant species in the North Carolina 1973 310 130 charter boat bottom fishing catches of 1974 597 59 reef fishes, accounting for 75% of the 1975 521 67 catch by number and 64% of the catch 1976 260 40 by weight (Manooch et al. 1981). The 1977 664 8 estimated recreational catch of black 1978 515 25 sea bass in the South Atlantic ranged 1979 624 100 from 3.6 million fish and 1.9 million 1980 694 97 kg in 1980 to 9.7 million fish and 1981 543 283 3.7 million kg in 1984 (Anonymous 1982 368 232 1984, 1985a,b, 1986 (Table 6). 1983 242 106 1984 449 72 Management

Mortality rates for black sea - = none reported. bass captured off South Carolina and Georgia from 1978 to 1981 were pre- sented by Low (1981) and Wenner et al. complete until age IV (Low 1981). The (1986). Catch curves for fish age at complete recruitment in deeper caught in traps at depths <40 m waters was about one year older. Low indicated that females were fully (1981) reported instantaneous rates recruited to the fishery at age 111, of total mortality that ranged from but the recruitment of males was not 0.75 to 0.83 for fish caught in water SEDAR25-RD15

Table 6. Recreational landings of mortality), black sea bass were sub- black sea bass for the South Atlantic jected to sufficient fishing mortal- region, 1980-1985 (Anonymous 1984, ity on the headboat grounds to account 1985a, b, 1986). for 80% to 98% of the maximal YPR. The South At1antic Fishery Management Counci 1 concluded from YPR analysis Year Number Weight (kg) that black sea bass are probably in (1000's) (1000's) the range of growth overfishing, i.e., excessive harvesting of small fish (Anonymous 1983). To control growth overf ishing and prevent recruitment overfishing (excessive harvesting of spawners which decreases recruitment), a minimum size 1imit of 203 mn (8 inches) and restrictions on fish trap dimensions and trap fishing areas were adopted for black sea bass caught in the South Atlantic Fishery Conservation Zone (Federal waters south of Cape Hatteras). Minimum mesh size of traps is 1 by 2 inches (rectangular) or 1 1/2 inches (hexagonal ) .

<40 m and from 0.60 to 0.61 for fish in deeper water. Wenner et al. (1986) ECOLOGICAL ROLE reported that the instantaneous rate of total mortality of age IV and older black sea bass from catch curve Black sea bass are carnivorous analysis, ranged from 0.72 in 1978 to bottom-feeders that eat crustaceans 1.32 in 1981 for comnercial fish (primarily crabs and shrimp), fish, traps and from 0.73 in 1979 to 1.43 mollus ks , and echinoderms (Smith 1907 ; in 1981 for hook and line. From Hildebrand and Schroeder 1928; Cupka 1978 to 1981, mortality rates in- et al. 1973; Link 1980; Steimle and creased from 51.3% to 73.3% for trap- Ogren 1982). Adults eat mainly crabs caught fish and from 51.6% to 76.1% and fish, and the young eat shrimp, for hook and line-caught fish older isopods, and amphipods. Link (1980) than age IV. found that the diet of black sea bass, based on frequency of occurrence, was Wenner et al. (1986) used composed primarily of crustaceans Petersen mark-recapture techniques to (91%). mollusks (28%), and fishes calculate population densities of 14 (28%). Crabs occurred in about 51% of to 125 individuals per hectare on the stomachs and shrimp in about 13%. two shallow-water patch reefs. Cupka et al. (1973) noted that the Abundance on both reefs declined from presence of barnacles and other ses- 1981 to 1983. Powles and Barans sile organisms such as colonial (1980) estimated black sea bass were indicative of some densities of 33 to 66 fish/ha and a grazing activity, particularly by biomass of 5.1 to 9.9 kg/ha from adults. Results of feeding studies are television transect data. Yield-per- affected by of collection gear-- recruit (YPR) models for black trawl -caught black sea bass had minimal sea bass presented by Huntsman et al. stomach eversion, but eversion was (1983) indicated that at the lowest significant in fish caught with hook- estimates of M (instantaneous natural and-line (Rogers et al. 1986). 9 SEDAR25-RD15

Fish assemblages associated with cardinalf ish (A o on seudomacul atus) hard bottom or reef habitats in the and jackknife-*f?o: is quetus South Atlantic Bight have been des- cribed in studies using submersibles, television and diver transects, trawls, and traps. Powles and Barans ENVIRONMENTAL REQUIREMENTS (1980) found that black sea bass, along with round scad (Decapterus and southern porgy Temperature !-As !-As aculeatus) , dominated the fish assemblage of two inshore Differences in temperature regime reef areas (16-32 m deep) off South appear responsible for differences in Carolina. Television estimates of seasonal distribution and migratory black sea bass and southern porgy tendency of black sea bass between the density and biomass were greater than South Atlantic and Mid-Atlantic Bights trawl estimates. Black sea bass (Musick and Mercer 1977). North of occurred singly, swimning near Cape Hatteras, North Carol ina, black bottom among attached organisms. sea bass move south and offshore in Chester et al. (1984) identified seven fall as inshore water temperatures groups of distributional ly similar decline. Black sea bass have been reef fish species on the continental caught at temperatures as low as 6 "C shelf of North Carolina and South in winter, but largest catches were Carolina, based on analyses of head- taken at bottom temperatures of 8-10 boat and research cruise catch re- "C and higher (Goode 1884; Nesbit and cords. Black sea bass was the domi- Neville 1935; Musick and Mercer 1977). nant member of an inshore comnunity on Tagging studies off the South Atlantic the inner shelf (<30 m) that was coast revealed that adult black sea also characterized by the presence of bass were year-round residents in whitebone porgy (Calamus leucosteus), given areas and, in contrast to spottai 1 porgy (msholbrooki), Mid-Atlantic sea bass, did not sheepshead porgy*mus penna), undertake seasonal migrations. The longspine porgy (Stenotomus ca rinus) lowest bottom temperatures off South tomtate (Haemul on ayrol ineahna Carolina in the depth range inhabited pigfish (Orthoistis chrysoptera) . by adult black sea bass were 10 "C Cluster ana+ ysis of summer and winter and higher during all seasons (Walford trawl collections in the South Atlan- and Wicklund 1968). Grimes et al. tic Bight grouped black sea bass with (1982) reported that a decrease in temperature at 1ive-bottom areas from 22 to 8 "C from November 1975 to February 1976 was accompanied by a (~onacanth;~ hispidus) , northern sea- marked change in faunal composition rob1n Prionotus carol inus), sand from a red porgy (Pa rus perch (I!? lectrum f-, inshore vermil ion snapper .-frriiomEY%L 1izardfi*dus foetens), and aurorubens)-whi te grunt -7I&iE scrawled cowf ish (~acto-quadri- assemblage to a= cornis), in summer-d+ Se erry and Van dominated by black sea bass, Atlantic -1984). Wenner ' s (1983) cluster spadefish (Chaetodi terus faber) , and analysis of a late spring sponge-coral tautog (Tautoga+ oni ti s habitat trawl survey also grouped black sea bass with cubbyu, northern In North Carol ina estuaries, searobin. sand ~erch,and in addition, black sea bass were collected at 6 to bank cusk-eel ' (0 hidion hol brooki ) , 29 "C (Link 1980; Schwartz et al. Atlantic nidshipiiichth s Plec- 1981). Cupka et al. (1973) reported ?trodon) bank seabasdrms that juvenile black sea bass were ocyurus) , whi tebone porgy, twospot taken in estuaries at water tempera- SEDAR25-RD15

tures of 5.6 to 30.4 "C, but were most (Link 1980) to depths of 73 to 165 m abundant in waters above 10 "C. offshore in winter north of Cape Hatteras, North Carol ina (Pearson Salinity 1932; Musick and Mercer 1977). Numer- ous studies of the South Atlantic Although the black sea bass is a Bight reef fish communities indicated marine species, juveniles and year- that black sea bass were most abundant 1ings inhabit high sal inity sections at depths of 16 to 38 m (Struhsaker of estuaries as nursery grounds. 1969; Powles and Barans 1980; Grimes In North Carol ina estuaries, black sea et al. 1982; Chester et al. 1984; bass were collected at salinities of Sedberry and Van Dolah 1984; Parker 1.0 to 36.0 ppt but most often at and Ross 1986). Black sea bass were >14.0 ppt (Link 1980; Schwartz et al. caught as deep as 46-51 m off a rocky 1981). Cupka et al. (1973) reported ledge on the Georgia coast in June that the most important sea bass 1986 (Wayne Waltz, S.C. Wildlife & nurseries were in high salinity 030 Marine Resources Dep. ; pers. comm. ). ppt) reef areas, and Dahlberg (1972) Sedberry and Van Dolah (1984) reported found juvenile sea bass mostly in the that they were more abundant in sumner lower reaches of a Georgia estuary. than in winter at inner shelf stations (16-22 m) , but seasonal differences in Substrate abundance were minor at middle shelf stations (25-38 m). The black sea bass is a dominant species associated with the hard- or live-bottom sponge-coral habitat in Contaminants the South Atlantic Bight (Struhsaker 1969; Powles and Barans 1980; Grimes Stone et al. (1975) used tires to et al. 1982; Wenner 1983; Chester et construct a simulated artificial tire al. 1984; Sedberry and Van Dolah 1984; reef to determine if pollutants would Parker and Ross 1986). This habitat, leach from the tires and affect black which is located on the open shelf sea bass. No significant increase in from depths of 18 to 55 m, consists of concentrations of zinc, organochlo- 1 ow rock ridges , outcroppi ngs , coral ride insecticides, or polychlorinated patches, and shipwrecks (Struhsaker biphenyls (PCB's) was found. Hall et 1969; Parker and Ross 1986). The a1 . (1978) determined the concentra- black sea bass is also one of the most tions of 15 trace elements in black abundant species on artificial reefs sea bass muscle and liver tissue, but along the South Atlantic coast did not report information on accept- (Buchanan 1973; Buchanan et al. 1974). able or safe levels. Fair and Fortner (1987) found that the hydrocarbon Depth benzo(a) pyrene (a compound found in petroleum) consumed by seabass Black sea bass occur at depths as increased muscle tissue concentrations shallow as 1 m in estuarine waters of cadmium fed to the fish. SEDAR25-RD15 SEDAR25-RD15

LITERATURE CITED

Anonymous. 1983. Fishery management Beaumariage, D.S. 1964. Results from plan, regulatory impact review, and the 1963 Schlitz tagging program. final environmental impact statement Fla. Board Conserv. Mar. Res. Lab. for the snapper-grouper fishery of Tech. Ser. 43. 34 pp. the South Atlantic region. South At1antic Fishery Management Counci 1 in coop. with National Marine Beaumariage, D.S. 1969. Returns from Fisheries Service. Charleston, S.C. the 1965 Schlitz tagging program. 89 pp + Appendices. Fla. Board Conserv. Mar. Res. Lab. Tech. Ser. 59. 38 pp.

Anonymous. 1984. Marine recreational fishery statistics survey, Atlantic Beaumariage, D.S., and A.C. Wittich. and gulf coasts, 1979 (Revised)- 1966. Returns from the 1964 Schlitz 1980. Natl. Mar. Fish. Serv. Curr. tagging program. Fla. Board Con- Fish. Stat. No. 8322. 239 pp. serv. Mar. Res. Lab. Tech. Ser. 47. 50 PP. Anonymous. 1985a. Marine recreation- al fishery statistics survey, At- Bortone, S.A. 1977. Osteological lantic and gulf coasts, 1981-1982. notes on the genus Centro ristis Natl. Mar. Fish. Serv. Curr. Fish. (Pisces : Serranidae) . Nort-T%m east u Stat. No. 8324. 215 pp. Sci. 1(1):23-33.

Buchanan, C.C. 1973. Effects of an Anonymous. 1985b. Marine recreation- artificial habitat on the marine al fishery statistics survey, At- sport fishery and economy of Mur- lantic and gulf coasts, 1983-1984. rells Inlet, South Carolina. U.S. Natl. Mar. Fish. Serv. Curr. Fish. Natl. Mar. Fish. Serv. Mar. Fish. Stat. No. 8326. 222 pp. Rev. 35 (9):15-22.

Buchanan, C.C., R.B. Stone, and R.O. Anonymous. 1986. Marine recreation- Parker, Jr. 1974. Effects of arti- al fishery statistics survey, At- ficial reefs on a marine sport lantic and gulf coasts, 1985. fishery off South Carolina. U.S. Natl. Mar. Fish. Serv. Curr. Fish. Natl. Mar. Fish. Serv. Mar. Fish. Stat. No. 8327. 130 pp. Rev. 36(11) :32-38.

Chester, A.J., G.R. Huntsman, P.A. Bearden, D.M., and M.D. McKenzie. Tester, and C. S. Manooch 111. 1971. An investigation of the off- 1984. South Atlantic Bight reef shore resources of fish comnunities as represented in South Carol ina. S.C. Wild1 . Resour. hook-and-1 ine catches. Bull. Mar. Dep. Tech. Rep. 2. 19 pp. Sci . 34 (2) :267-279. SEDAR25-RD15

Cupka, D.M., R.K. Dias, and J. Tucker. Hardy, H.D., Jr. 1978. Serranidae- 1973. Biology of the black sea sea basses. Pages 35-68 in Develop- bass, Centro Fistis striata (Pisces: ment of fishes of the ~m-Atlantic Serran+ dae from South Carolina Bight. An atlas of egg, larval and waters. S:C. ~ildl. Mar. ~esour. juvenile stages. Vol. 3. Aphredo- Dep. Unpubl . MS. 93 pp. deridae through Rachycentridae. U.S. Fish Wildl. Serv. Biol. Serv. Program FWS/OBS-78/12. 394 pp. Dahlberg, M.D. 1972. An ecological study of Georgia coastal fishes. Hildebrand, S.F., and W.C. Schroeder. U.S. Natl. Mar. Fish. Serv. Fish. 1928. Fishes of Chesapeake Bay. Bull. 70(2) :323-353. U.S. Bur. Fish. Bull. 43. 366 pp. Hoff, F.H., Jr. 1970. Artificial Fair, P.H., and A.R. Fortner. 1987. spawning of black sea bass, Centro- Effect of ingested benzo(a) pyrene ristes striatus me1 anus insb burg, and cadmi um on tissue accumulation, kby-nic gonadotrophic hydroxyl ase activity, and intestinal hormones. Fla. Dep. Nat. Resour. morphology of the black sea bass, Mar. Res. Lab. Spec. Scl. Rep. Centro~ristisstriata. Environ. Res. No. 25. 17pp. 42:185-195. Huntsman, G.R. 1976. Offshore head- boat fishing in North and South Fischer,W.,ed. 1978. FA0 species Carolina. U.S. Natl. Mar. Fish identification sheets for fishery Serv. Mar. Fish. Rev. 38(3):13-23, purposes. Western Central At1 antic. FAO, Rome, Vol. 4. Huntsman, G.R., and C.S. Manooch 111. 1978. Coastal pelagic and reef fishes in the South Atlantic Bight. Frame, D.W., and S.A. Pearce. 1973. Pages 97-106 in R.H. Stroud, ed. A survey of the sea bass fishery. Marine recreatGnal fisheries 3. U.S. Natl. Mar. Fish. Serv. Mar. Sport Fishing Institute., Wash- Fish. Rev. 35(1-2) :19-26. ington, D.C. 176 pp.

Goode, G.B. 1884. The fisheries and Huntsman, G.R., C.S. Manooch 111, and fishery industries of the United C.B. Grimes. 1983. Yield per re- States. Sect. I. Natural history cruit models of some reef fishes of of useful aquatic . 895 pp. the U.S. South Atlantic Bight. U.S. Natl. Mar. Fish. Serv. Fish. Bull. 81(4) :679-695. Grimes, C.B., C.S. Manooch, and G.R. Huntsman. 1982. Reef and rock Kendal 1 . A.W., Jr. 1972. DeScri~tion outcropping fishes of the outer of black sea bass, continental shelf of North Carolina striata (Linnaeus), and South Carolina, and ecological occurrences north of Cape Lookout. notes on the red porgy and vermilion North Carolina, in 1966. ' U.S. NatlI snapper. Bull. Mar. Sci . 32(1) :227- Mar. Fish. Serv. Fish. Bull. 70(4): 289. 1243-1260. Hall, R.A., E.G. Zook, and G.M. Mea- Kendall , A.W., Jr. 1977. Biological burn. 1978. National Marine Fish- and fisheries data on black sea eries Service survey of trace ele- bass, Centro ristis striata (Lin- ments in the fishery resource. naeus) .*. TZK ~erv.. U.S. Natl. Mar. Fish. Serv. Spec. NEFC Sandy Hook Lab., Tech. Ser. Sci. Rep. Fish. 721. 313 pp. Rep. No. 7. 29 pp. SEDAR25-RD15

Lavenda, N. 1949. Sexual differences Bur. Fish. Fish. Circ. No. 18. and normal protogynous hermaphrodi t- 12 PP. ism in the Atlantic sea bass, Cen- tro ristes striata. Copeia 1949 *4. *4. Parker, R.O., Jr., and S.W. Ross. 1986. Observing reef fishes from Link, G.W., Jr. 1980. Age, growth, submersibles off North Carol ina. reproduction, feeding and ecological Northeast Gulf Sci . 8(1) :31-49. observations on the three s~eciesof Centro ristis (Pisces : ~erranidae) Pearson, J .C. 1932. Winter trawl 777dkErolina waters. Ph.D. fishery off the Virginia and North Thesis. University of North Caro- Carolina coasts. U.S. Bur. Fish. lina, Chapel Hill. 277 pp. Invest. Rep. No. 10. 31 pp.

Low, R.A., Jr. 1981. Mortality rates Powles, H., and C.A. Barans. 1980. and management strategies for black Groundfish moni toring in sponge- sea bass off the southeast coast of coral areas off the Southeastern the United States. N. Am. J. Fish. United States. U.S. Natl. Mar. Manage. l(2) :93-103. Fish. Serv. Mar. Fish. Rev. 42(5): 21-35. Manooch, C.S., 111, L.E. Abbas, and J.L. Ross. 1981. A biological and Reinboth, R. 1965. Sex reversal in economic analysis of the North Caro- the black sea bass Centro ristes lina charter boat fishery. U.S. striatus. Anat. RecA Natl. Mar. Fish. Serv. Mar. Fish. m. Rev. 43(8) :1-11. Rivers, J.B. 1966. Gear and tech- Mercer, L.P. 1978. The reproductive nique of the sea bass trap fishery bioloov and DODU~ation dynamics of in the Carolinas. Comer. Fish. black-" sea bass; Centro ristis c- Rev. 28(4) :15-20. ata. Ph.D. Thesheaeof mliam and Mary, ~illiamsbur~;Va. 196 pp. Robins, C.R., R.M. Bailey, C.E. Bond, J.R. Brooker, E.A. Lachner, R.N. Miller, R.J. 1959. A review of the Lea, and W.B. Scott. 1980. A list sea basses of the genus Centro- of common and scientific names of ristes (Serranidae) . Tulane Stud. fishes from the United States and h(2) :35-68. Canada. Am. Fish. Soc. Spec. Sci. Publ. No 12. 174 pp. Moe, M.A., Jr. 1966. Tagging fishes in Florida offshore waters. Fla. Board Conserv. Mar. Lab. Tech. Ser. Rogers, S.G., H.T. Langston, and T.E. No. 49. 40 pp. Targett . 1986. Anatomical trauma to sponge-coral reef fishes captured by Musick, J.A., and L.P. Mercer. 1977. trawling and angling. U.S. Natl. Seasonal distribution of black sea Mar. Fish. Serv. Fish. Bull. 84:697- bass, Centro ristis striata, in the 704. Mid-AtlanticA Big t withmments on the ecology and fisheries of the species. Trans. Am. Fish. Soc. 106 Schwartz, F.J., W.T. Hogarth, and M.P. (1) :12-25. Weinstein. 1981. Marine and fresh- water fishes of the Cape Fear Nesbit, R.A., and W.C. Neville. 1935. Estuary, North Carol ina, and their Conditions affecting the southern distribution in relation to environ- winter trawl fishery. Bull. U.S. mental factors. Brimleyana 7:17-37. SEDAR25-RD15

Sedberry, G.R., and R.F. Van Dolah. Board Conserv. Mar. Res. Lab. Prof. 1984. Demersal fish assemblages Pap. Ser. No. 5. 76 pp. associated with hard bottom habitat in the South Atlantic Bight of the Walford, L.A., and R.I. Wicklund. U.S.A. Environ. Biol. Fish. ll(4): 1968. Monthly sea temperature 241-258. structure from the Florida Keys to Cape Cod. Serial Atlas of the Marine Environment . Folio 15. Amer- Smith, H.M. 1907. The fishes of North ican Geographical Society, N.Y. Carolina. N.C. Geol. Econ. Surv. 11. 423 pp. Waltz, W., W.A. Roumillat, and P.K. Steimle, F.W., Jr., and L. Ogren. Ashe. 1979. Distribution, age 1982. Food of fish collected on structure, and sex composition of artificial reefs in the New York the black sea bass, Centro ristis Bight and off Charleston, South striata, sampled alongli--G t e sout Carolina. U.S. Natl. Mar. Fish. eastern coast of the United States. Serv. Mar. Fish. Rev. 44(6-7) :49-52. S.C. Mar. Resour. Cent. Tech. Rep. No. 43. 18pp. Stone, R.B., L.C. Coston, D.E. Hoss, and F.A. Cross. 1975. Experiments on some possible effects of tire Wenner, C.A. 1983. Species associa- reefs on pinfish (La odon G- tions and day-night variability of boides) and black sea% ass (Centro- trawl-caught fishes from the inshore ristis striata). U.S. ~atl.. sponge-coral habitat, South At1 antic bServ. Mar. Fish. Rev. 37(3): Bight. U.S. Natl. Mar. Fish. Serv. 18-20. Fish. Bull. 81(3) :537-552.

Struhsaker, P. 1969. Demersal fish resources: composition, distri bu- Wenner, C.A., W.A. Roumillat, and C.W. tion, and commercial potential of Waltz. 1986. Contributions to the the continental shelf stocks off life history of black sea bass. Southeastern United States. U.S. ~entropristis striata, off the Fish Wildl. Serv. Fish. Ind. Res. Southeastern United States. U.S. 4(7) :261-300. Natl. Mar. Fish. Serv. ~ish.~ull. 84(3) :723-741. Tagatz, M.E. 1967. Fishes of the St. Johns River, Florida. Q. J. Fla. Acad. Sci . 30(1) :25-59. Wilson. H.V. 1891. The embrvoloav of the -sea bass (Serranus Grariis). Topp, R. 1963. The tagging of fishes U.S. Fish Comm.m -9. in Florida, 1962 program. Fla. SEDAR25-RD15

50272 -101 REPORT DOCUMENTATION 2. 3. Recipient's Accession NO. PAGE Bioloctical Rewrt 82(11.99)* I - 1I 4. Title and Subtitle Species Profiles: Life Hi stories and Environmental Requirements of Coastal Fishes and Invertebrates (South Atlantic)--Black Sea

7. Author(s) Linda P. Wrcer 9. Performing Organization Name and Addrass 10. Pmied/Task/Worll Unit No.

111. contract(c) or Grant(C) No. I

-. 12. Sponsoring Organization Name and Addnss

U.S. Department of the Interior U.S. Army Corps of Engineers 13. TYW of Report F'mriod Cownd Fish and Wildlife Service Waterways Experiment Station Research and Development P.O. Box 631 7 National Wetlands Research Center Vicksburg, MS 39180 Washington, DC 20240

IS. Supplamentary Notes 1 *U.S. Army Corps of Engineers Report No. TR EL-82-4 16. Abstract (Limit: 2W word.) Species profiles are literature summaries on the taxonomy, morphology, range, life history, and environmental requirements of coastal aquatic species. They are designed to assist in environmental inpact assesment. The black sea bass, striata, is an abundant species associated with the inshore sponge-coralXXTat in the South Atlantic Bight (Cape Hatteras to Cape Canaveral). It is a protogynous hermaphrodite (each individual is first a female and then a male) that spawns from January to June on the Continental Shelf. Juveniles utilize estuaries, as well as offshore areas, for nurseries. It is a slow grwing species with a life span of about 10 years. Juveniles and adults are bottom-feeding carnivores. Adults have been collected at temperatures as low as 6 "C but are most abundant at temperatures of 8 to 10 "C and above. Juveniles tolerate lower tenperatures and greater salinity ranges than adults. Black sea bass are primarily harvested by the recreational hook and line fishery and the commercial trap fishery. Yield-per-recruit analyses indicate that the harvest of black sea bass is less than the maximum possible due to a conbination of high fishing pressure and harvest of small fish.

17. Document Analysis a. Deuripton Estuaries Tenperature Fi shes Sediments Fisheries Feeding habits Sal ini ty Life cycles Growth Contaminants b. Identifien/~n-EndedTerms

Black sea bass Environmental requirements Centropri stis striata

18. Avallablllty Statament 19. Scurity Class (This Report) 21. No. of Pages Unclassified 16 Unl imi ted 9. price Unclassified (Sae ANSI-239.18) OPTIONAL FORM 272 (4-77) SEDAR25-RD15

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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