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American Shad Alosa sapidissima

Atlantic Croaker Micropogonias undulatus

Bay Anchovy Anchoa mitchilli

Black Drum Pogonias cromis

Flathead Catfish Pylodictis olivaris

Hardhead Catfish Arius felis

Largemouth Bass Micropterus salmoides

Mummichog Fundulus heteroclitus

Red Drum Sciaenops ocellatus

Redbreast Sunfish Lepomis auritus

Sheepshead Archosargus probatocephalus

Shortnose Sturgeon Acipenser brevirostrum

Silver Perch Bairdiella chrysoura

Southern Flounder Paralichthys lethostigma

Southern Kingfish Menticirrhus americanus

Spot Leiostomus xanthurus

Spotted Seatrout Cynoscion nebulosus

Striped Bass Morone saxatilis

Striped Mullet Mugil cephalus

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American Shad Alosa sapidissima

American shad

Description

American shad are the largest members of the family Clupeidae (which includes herrings, alewives, menhaden and sardines) in the United States. Clupeids are generally deep-bodied, silvery fishes that have relatively large, easily-shed scales and often associate in large schools. They have rough scales, or scutes, along the ventral edge of the body, forming a "saw-belly," a single dorsal fin midway along the body, and a deeply forked caudal fin. In addition, Clupeids have no lateral line and lack spines on all their fins. American shad are a metallic green color dorsally with silvery sides and have a dark shoulder spot behind the gill cover, sometimes followed by a series of smaller spots.

Habitat and Biology

American shad are distributed from the Gulf of St. Lawrence to Florida and are most abundant between Connecticut and North Carolina. They are anadromous; that is, they migrate upriver from brackish water to spawn. Upstream migration coincides with favorable water temperatures and typically begins in the spring. In South Carolina, adult migration to freshwater begins in January, peaks in February and March, and ends in April. Similar to salmon, American shad return to the same tributary system where they were hatched.

Spawning usually occurs in tidal and nontidal freshwater areas over shallow flats, sometimes hundreds of miles inland, where currents are strong enough to keep the eggs suspended in the water. Usually, American shad travel far enough upstream that the drifting eggs hatch before reaching saltwater. Males are 3-4 years old and females are 4-5 years old at sexual maturity. They spawn only once or twice (rare in South Carolina) during their life. During spawning, females are followed by 1-3 males who fertilize the eggs as they are released into the water. Each female may release between 100,000 and 700,000 eggs over the spawning season. The eggs sink slowly and drift along the bottom of the river, hatching within 4-6 days. Larval and juvenile shad move downriver towards the ocean but remain in the rivers and estuaries until the following spring, when they move into the ocean and join the adult population. They remain there in large schools during fall and early winter. However, not all adults survive the spawning event. In northern latitudes, where water temperature changes gradually and shad do not migrate far upstream, large numbers of adults find their way back to the ocean and survive to spawn the following year. In the southern part of their range, however, water warms up rapidly in the spring and shad tend to move well upriver to find suitable spawning sites. Consequently, less than 10% of spawning adults survive the migration back out to sea. Juveniles spend an average of four years along the Atlantic coast before they undertake their first spawning migration. This species lives to be 5 to 7 years old.

American shad are primarily planktivores. Juvenile shad feed on small invertebrates, insects, fish eggs, and algae. Adults use their gill rakers (comb-like structures on the gill arches) to filter small planktonic animals from the water for food during the riverine and oceanic phases of their life cycle. Adults do not feed during the upstream migration to their spawning areas.

Species Significance

American shad are important commercial and game fish. Fisheries for American shad have existed along the Atlantic coast since the nineteenth century, but have been declining since then due to overfishing and degradation of spawning habitat. In South Carolina, the shad fishery is the single most important finfish fishery. Since 1987, there has been a continuous decline in numbers of shad landed; however, landings in 1996 were considerably higher than in previous years. Female shad, referred to as roe shad, are prized for their egg mass (roe), which is considered a delicacy. Male shad, called bucks, are popular for their meat. In South Carolina, a recreational fishery for American shad exists exclusively in fresh water where fish are taken with dip nets or with artificial lures during their spawning migration.

The once huge populations of shad started to decline in the mid nineteenth century due to dam construction, pollution, and overfishing. However, stocks have shown some recovery in recent years. In the Edisto, Ashepoo and Combahee Rivers, however, shad populations are not as healthy as they were in years past. This species is currently not threatened or endangered.

References

MacKenzie, C., L.S. Weiss-Glanz, and J.R. Moring 1985. Species profiles: Life histories and environmental requirements of coastal fishes and invertebrates (Mid-Atlantic): American shad. Biological Report 82(11.37). U.S. Department of Interior, Fish and Wildlife Service, Division of Biological Services, Washington, DC.

Moore, C. 1996. Review of the 1995 finfish season. South Carolina Department of Natural Resources, Charleston, SC.

Murdy, E.O., R.S. Birdsong, and J.A. Musick. 1997. Fishes of the Chesapeake Bay. Smithsonian Institution Press, Washington, DC.

Walburg, C.H. and P.R. Nichols. 1967. Biology and management of the American shad and status of the fisheries, Atlantic coast of the United States, 1960. Special Scientific Report - Fisheries No. 550. U.S. Department of Interior, Fish and Wildlife Service, Washington, DC.



Atlantic Croaker Micropogonias undulatus

Atlantic croaker

Description

As members of the drum family (Sciaenidae), croaker are well known for producing "drumming" sounds, which they do by vibrating special muscles on either side of their swimbladder. They are common inhabitants of South Carolina estuaries, including the ACE Basin, during spring and summer months. Croaker are easily recognized by numerous short barbels on either side below the mouth and a sharply jagged preopercle. The latter can be painfully evident upon holding a croaker, as the fish tend to flare their opercles when disturbed or frightened. Croaker have a rather long head with an inferior mouth, appropriate for feeding on the bottom. They have silvery bodies with a series of copperish or brownish markings arranged in diagonal bars that tend to become less distinct as the fish grows. Pectoral and pelvic fins are bright yellow to orange.

Habitat and Biology

Atlantic croaker are in the same family as spot (Sciaenidae) and have a life history similar to spot. This species has a protracted spawning season from late summer through early spring; however, peak reproductive activity is in late fall. Spawning takes place in near-shore ocean waters, and the resulting early-life stages utilize some of the same behavioral patterns as spot to gain entrance to estuarine systems. Atlantic croaker are about 1/4 inch (4-6 mm) at the time of recruitment into the estuarine habitats. This species also utilizes the shallow marsh habitat as its primary nursery area. The small fishes feed on crustaceans and benthic infauna and epifauna. After a short residence period, they move from the shallows to other areas of the estuary such as the channels. Some Atlantic croakers are sexually mature between age one and two, and all are mature the following year. Throughout their life, Atlantic croakers eat small crustaceans and small fishes either on the bottom or near the bottom.

Species Significance

Landings of Atlantic croaker in South Carolina have represented a limited contribution to the commercial catch for the South Atlantic region. Since 1950, commercial landings in South Carolina have fluctuated widely, reaching a peak in the early 1970s. As of 1987, when a Fishery Management Plan for Atlantic croaker was instituted by the Atlantic States Marine Fisheries Commission (ASMFS), commercial landings in South Carolina resulted mainly from a limited fall haul seine fishery and from shrimp trawl bycatch. Currently, most Atlantic croaker that are sold commercially are caught incidentally.

The recreational harvest in South Carolina is also relatively limited. Catches since 1981 were at their highest in 1984 and at their lowest in 1993. Currently, there are no restrictions governing the catch of Atlantic croaker in South Carolina.

References

McErlean, A.J., S.G. O'Connor, J.A. Milhursky, and C.I. Gibson. 1973. Abundance, diversity and seasonal patterns of estuarine fish populations. Estuarine Coastal Marine Science 1:19-36.

Miglarese, J.V., C.W. McMillan, and M.H. Shealy, Jr. 1982. Seasonal abundance of Atlantic croaker (Micropogonias undulatus) in relation to bottom salinity and temperature in South Carolina estuaries. Estuaries 5(3):216-223.

Moore, C. 1993. South Carolina report for Atlantic croaker. In: Proceedings of a Workshop on spot, (Leiostomus xanthurus) and Atlantic croaker (Micropogonias undulatus). Special Report No. 25. Atlantic States Marine Fisheries Commission, Washington, DC.

Wenner, C. 1998. Personal communication. South Carolina Department of Natural Resources, Marine Resources Division, Charleston, S.C.



Bay Anchovy Anchoa mitchilli

bay anchovy

Description

The bay anchovy is one of 10 species of anchovies on the Atlantic coast of North America and is the most abundant fish in estuarine and coastal habitats along the eastern United States. Bay anchovies (and other anchovy species) are similar in appearance to fishes of the herring family (Clupeidae). However, they can be distinguished by a prominent silver stripe on either side of the body and lack of scutes (bony scales) along their bellies. Bay anchovies are of a greenish color above and silvery below and have a single dorsal fin, which is located midway along the body. They are often confused with silversides (Menidia spp.), but the two can be easily distinguished--anchovies lack a spine in the dorsal fin and have a large, gaping mouth that extends almost to the edge of the opercle, whereas silversides have two distinct dorsal fins, the first with four spines, and a very small mouth that is tilted upwards.

Habitat and Biology

Along the Atlantic coast, the species is distributed from the Gulf of Maine to Florida. It is also abundant along the Gulf coast to Yucatan, Mexico. Bay anchovies inhabit primarily shallow bays and estuaries but also occur in tidal freshwater habitats.

Bay anchovies, as other members of the family Engraulidae, typically aggregate in large schools. They are planktivorous fish which use gill rakers--comb-like structures on their gill arches--to strain the water for food. In South Carolina, spawning occurs in the evening during the summer months. Eggs are pelagic, and larvae hatch within 24 hours. Growth in this species is rapid, especially at higher temperatures, with a fish reaching maturity a few months after hatching. Bay anchovies seldom live past the age of two.

Species Significance

Bay anchovies are often very abundant and are important food for commercially valuable species such as striped bass, bluefish, spotted seatrout and southern flounder. They also are a very important link between the plankton community and higher order consumers. Thus, even though the species itself has no recreational or commercial value, it fulfills a crucial role in the coastal food web.

References

Dorsey, S.E., E.D. Houde, and J.C. Gamble. 1996. Cohort abundances and daily variability in mortality of eggs and yolk-sac larvae of bay anchovy, Anchoa mitchilli, in Chesapeake Bay. Fishery Bulletin 98:257-267.

Jones, P.W., F.D. Martin, and J.D. Hardy, Jr. 1978. Development of fishes of the mid-Atlantic Bight: an atlas of egg, larval, and juvenile stages. Volume I: Acipenseridae through Ictaluridae. U.S. Fish and Wildlife Service, Office of Biological Programs. FSW/OBS-78/12. Ft. Collins, CO.

Luo, J. and J.A. Musik. 1991. Reproductive biology of the bay anchovy in Chesapeake Bay. Transactions of the American Fisheries Society 120:701-710.

Murdy, E.O., R.S. Birdsong, and J.A. Musick. 1997. Fishes of the Chesapeake Bay. Smithsonian Institution Press, Washington, DC.



Black Drum Pogonias cromis

black drum

SoundSound, SCDNR

Description

Black drum are large, long-lived members of the drum family (Sciaenidae), reaching over 1 meter (3.3 ft) in length and weighing over 50 kg (110 lbs). They are one of only four members of this family that possess chin barbels set in pairs. Along the South Carolina coast, only the Atlantic croaker, Micropogonias undulatus, shares this characteristic. Black drum, as their name suggests, are dark in color with a silvery luster. Their body shape is characteristic in that their back is distinctly elevated. When young, they possess four to five dark vertical bars that fade and eventually disappear as the animal grows.

Habitat and Biology

Black drum are members of the family Sciaenidae, and they spawn during the spring (late March through April) in high salinity waters near inlets. Eggs and larvae are planktonic, and eventually the larvae gain access to the shallow dendritic creeks of the tidal marsh habitat. Juveniles remain in these creeks for four to six months, after which they are found on the shallow tidal mud flats, often co-occurring with red drum and southern flounder. Black drum feed on invertebrates such as small clams and crustaceans, which are easily crushed by the strong molar-shaped pharyngeal teeth. These prey items are commonly found in association with oyster bars, and catches of small black drum (150-500 mm) occur primarily in these shallow-water habitats.

Black drum have a long life span, with the oldest fish being 30+ years of age. They have a fast growth rate for the first three years, after which the rate slows considerably. Sexual maturity appears to be attained at age 3 to 5. At the present time, there are numerous gaps in the life history of this species.

Species Significance

Historically, black drum were harvested commercially in South Carolina. Trotlines were set in the spring and baited with crabs to catch the large fish entering the estuaries. Currently, no commercial landings are reported from South Carolina, and the National Marine Fisheries Service offers no data on commercial landings for any state in the South Atlantic. Black drum are caught recreationally in South Carolina with hook and line in deep holes near the mouth of estuaries. There are no size restrictions or bag limits governing the recreational harvest of this species.

References

McErlean, A.J., S.G. O'Connor, J.A. Milhursky, and C.I. Gibson. 1973. Abundance, diversity and seasonal patterns of estuarine fish populations. Estuarine Coastal Marine Science 1:19-36.

Murdy, E.O., R.S. Birdsong, and J.A. Musick. 1997. Fishes of the Chesapeake Bay. Smithsonian Institution Press, Washington, DC.

Roumillat, W. 1998. Personal communication. South Carolina Department of Natural Resources, Marine Resources Division, Charleston, S.C.

Wenner, C. 1998. Personal communication. South Carolina Department of Natural Resources, Marine Resources Division, Charleston, S.C.



Flathead Catfish Pylodictis olivaris

flathead catfish

Description

Catfishes, members of the family Ictaluridae, are easily recognized by conspicuous barbels around their mouths and scaleless bodies. The pectoral and dorsal fins are equipped with spines that can inflict wounds causing considerable discomfort, depending on the particular species. The flathead catfish is the only member of the genus Pylodictis. This species can be distinguished by a white blotch on the dorsal lobe of the caudal fin (except in large adults), an adipose fin which is fused to the body along its entire length, and an anal fin with less than 17 rays. The fish are typically yellowish with brown mottling. The species gets its scientific name from its flattened head, protruding lower jaw, and coloration. Adults commonly reach 115 cm (45 in) long and generally weigh up to 23 kg (50 lbs). The current South Carolina record weighed 33.5 kg (74 lbs).

Habitat and Biology

Flathead catfish are typically found in freshwater rivers and reservoirs, but they can also inhabit brackish water. They are benthic inhabitants and utilize their barbels to locate food in the substrate. They are very secretive and spend the daylight hours associated with some type of aquatic cover. Foraging occurs at night, often in areas so shallow that their dorsal fins are exposed. Juveniles feed on immature insect larvae in riffle areas, and adults primarily target fish and crayfish. Adults are generally a solitary species, except during the spawning season. They prefer deep pools with heavy submerged cover. At night they move to shallow areas to feed. Young fish can be found in shallower areas around roots or riprap.

This species spawns in June or July. The eggs are laid in a large nest fanned out on the bottom in a natural cavity or near a large, submerged structure. An egg mass of up to 100,000 eggs can be produced by a large female. The eggs are agitated continuously by the male parent, which helps to provide oxygen and flush away silt from the nest. After the eggs hatch, the fry remain in a compact school around the nest for several days, guarded by the male fish. They soon disperse and begin a solitary existence. The fry first feed on insect larvae, but as they grow, fish and crayfish become more important in the diet. Large adults are almost completely piscivorous. The species reaches sexual maturity at 4 or 5 years of age. Trophy size can be reached in about 10 years.

Species Significance

Flathead catfish are not native to the ACE Basin but are originally from the Mississippi River drainage. They were first introduced into Lake Thurmond and Lake Marion in the early 1960s. North Carolina stocked the species in the Pee Dee drainage, and they have migrated from there downstream into South Carolina. Because of unauthorized releases, they are now found in several coastal rivers of the state, including the Edisto River in the ACE Basin. In the Edisto River, they have been found throughout the main branch and are beginning to move into the north and south forks.

In the Edisto River system, flathead catfish have had profound effects on native catfish and sunfish species. All three of the once plentiful bullhead catfish species are now rare. Redbreast sunfish populations appear to have also been drastically reduced since the introduction of flathead catfish into the system. Because flathead catfish have been found in estuarine reaches of the river, it is speculated that they may also impact some marine and amphidromous species as well. Most notable would be their potential impacts on shortnose sturgeon, which use the estuarine reaches of the river as feeding grounds during the winter. Currently, the Freshwater Fisheries Section of the South Carolina Department of Natural Resources is gathering data on life history information, and investigating the impacts of this species on the ecology of the Edisto River.

This species is important commercially and is a popular game fish in many areas. (See related section: Recreational Fisheries.)

References

Allen, D. 1998. pers. comm. South Carolina Department of Natural Resources, Freshwater Fisheries Division, Barnwell, SC.

Allen, D.E. and C.S. Thomason. 1993. Survey and inventory of fisheries resources in the Combahee River. Study Completion Report F-32. SC Wildlife and Marine Resources Department, Columbia, SC.

Thomason, C.S., D.E. Allen, and J.S. Crane. 1993. A fisheries study of the Edisto River, SC. Wildlife and Marine Resource Department, Study Completion Report, Federal Aid Projects F-32 and F-30.



Hardhead Catfish Arius felis

hardhead catfish

Description

Hardhead catfish are members of the family Ariidae. Even though members of this family are commonly known as sea catfishes, they are not confined to marine waters. Sea catfishes can be differentiated from freshwater catfishes (family Ictaluridae) by the number of barbels. Sea catfishes have three pairs, whereas freshwater catfishes have four. As other catfishes, hardhead catfish lack scales and they possess a fleshy adipose fin. Hardhead catfish vary in color, from dark brown to dark blue and reach up to 40+ cm (16 in) in length.

Habitat and Biology

Hardhead catfish are a seasonal resident of the ACE Basin. They first appear as water temperatures moderate in the late spring, and they remain in the system until the water cools in the fall. The catfish then move off-shore and south, where they over-winter. Hardhead catfish spawn during the early summer in estuarine and near-shore waters along the South Carolina coast. The large (8-12 mm) fertilized eggs are collected by the male, and held in his mouth until hatching. Males do not feed during the one-month period while larvae and small juveniles are protected in this fashion. Little is known about growth rates or age at maturity for this species. The only obvious seasonal migration is from shallow estuarine areas during cold water temperatures to warmer waters in lower latitudes. Hardhead catfish feed on epibenthic crabs, shrimps and small fishes.

Species Significance

Hardhead catfish are of no commercial value, and they represent a very small portion of the recreational catch in South Carolina. This species is abundant in South Carolina waters.

References

Jones, P.W., F.D. Martin, and J.D. Hardy, Jr. 1978. Development of fishes of the mid-Atlantic Bight: an atlas of egg, larval, and juvenile stages. Volume I: Acipenseridae through Ictaluridae. U.S. Fish and Wildlife Service, Office of Biological Programs. FSW/OBS-78/12. Ft. Collins, CO.

Murdy, E.O., R.S. Birdsong, and J.A. Musick. 1997. Fishes of the Chesapeake Bay. Smithsonian Institution Press, Washington, DC.

Roumillat, W. 1998. Personal communication. South Carolina Department of Natural Resources, Marine Resources Division, Charleston, S.C.



Largemouth Bass Micropterus salmoides

largemouth bass

Description

The genus Micropterus includes some of the well-known freshwater game fishes including the small- and largemouth basses. The latter belong to the sunfish family (Centrarchidae) and should not be confused with either the temperate basses of the family Percichthyidae, which includes the striped and white basses, or the sea basses and groupers of the family Serranidae. They are the largest members of the sunfish family, often reaching a weight of 2-4 kg (5-8 lbs), and the South Carolina record is currently over 7 kg (16 lbs). Large and smallmouth bass can be distinguished from other sunfish in the family Centrarchidae by their rather elongate body and olive coloration.As with all sunfish, largemouth bass have a spinous first dorsal fin followed by a soft-rayed second dorsal. Coloration depends on water clarity, but generally their color is olivaceous, green dorsally fading to a pale underside with a distinct black stripe running from the fish's opercular flap to the base of the tail. The corner of the mouth extends past the eye (thus the name "large mouth"), and no teeth are present on the fish's tongue. The fish has a moderately elongate and slightly compressed body. They usually have 12 dorsal rays and 3 anal spines.

Habitat and Biology

The native range of largemouth bass is from the lower Great Lakes to the Mississippi River drainage, the Gulf Coast, and Florida. Along the Atlantic coast, the species occupies rivers and ponds as far north as Virginia. Bass are generally confined to fresh water, but they may move into low salinity areas of tidal rivers for short periods of time. They prefer very slow-moving water, especially lakes and larger streams or rivers, inshore waters of ponds, lakes, reservoirs, sloughs of the Delta, creeks, and some irrigation ditches. Largemouth bass are often found around or near submerged structures such as a fallen trees, stumps, rocks, grasses, or an overhanging bank.

The largemouth bass is native to the ACE Basin. In the ACE Basin the largemouth bass is commonly found in the riverine ecosystem, a freshwater community. They prefer slow-moving water and are found throughout the area from small creeks to the low salinity areas of the estuary. They are often stocked in private impoundments as a predatory species. They commonly lurk near cover during the day and move into shallow areas to feed at dark.

Largemouth bass found in the ACE Basin appear to be closely linked genetically to the Florida subspecies. Sexual maturity occurs in the second year. Spawning occurs in the spring from late April to June when water temperatures near 65°F. Spawning occurs over a nest constructed along the shallow margins of their habitat. Male fish clear a shallow, circular depression in a gravelly or sandy bottom of a deep pool and establish a territory. Several females spawn in the nest of a single male. The female deposits up to 1 million eggs that are fertilized by the male as they are deposited in the nest. The male then devotes full time to guarding the nest from predators and fans the eggs to increase oxygen levels and reduce siltation on the nest. While guarding the nest and the young, he does not feed. When the fry hatch they form a school and stay around the nest for several days. The attentive male continues to protect the fry for some time after they leave the nest.

Young bass feed on zooplankton, insects, and small fishes and are also sometimes cannibalistic. They grow up to 20 cm (8 in) in the first year. Adults are top level carnivores in their respective habitats, feeding primarily on crawfish and many fish species, as well as on snakes, frogs, and even baby ducks. Feeding activity takes place during both day and night and is initiated by both hunger and reflex. Largemouth bass are sight feeders and actively hunt for food. This genetic reflex is behind the success of many of the fishing lures used to capture them. The average maximum life span of largemouth bass is typically 10 to12 years.

Species Significance

Along with trout, largemouth bass are the most sought after game fish in the United States. The world record largemouth was 10 kg (22.25 lbs.) captured in Montgomery Lake, Georgia. In South Carolina, fishing for largemouth bass occurs in lakes and creeks in riverine ecosystems.

References

Chew. R.L. 1968. Annual progress report for investigations project: Largemouth bass study. State of Florida Game and Freshwater Fish Commission, Tallahassee, FL.

Thomason, C.S., D.E. Allen, and J.S. Crane. 1993. A fisheries study of the Edisto River, SC. Wildlife and Marine Resource Department, Study Completion Report, Federal Aid Projects F-32 and F-30.

Wisconsin Conservation Department. 1961. The largemouth bass: Its life history, ecology, and management. Publication 232-67. Madison, WI.



Mummichog Fundulus heteroclitus

Mummichog

Description

Mummichogs, locally called mud minnows, belong to a group of fishes known collectively as killifish. Killifish are members of the family Cyprinodontidae. They make up a large portion of the small fish in tidal creeks, saltwater marshes, lagoons, and other shallow coastal habitats. Mummichogs have a single, spineless dorsal fin and pelvic fins positioned near the anal fin. The head is flattened and the mouth is tilted upwards. Most species in the family Cyprinodontidae are sexually dimorphic; that is, males and females look different. Small female mummichogs are brownish above and pale below with 12-15 dark vertical bars. The dorsal and anal fins are tinged with green. Distinctive markings disappear as females grow larger. Males are darker in color, usually dark green or olive with a yellowish underside; spawning males have a dark spot towards the rear of the dorsal fin. Their sides are silvery with approximately 15 vertical bars and many small spots that continue on to the f in membrane, varying in color from white to yellow. Maximum adult size is 12 cm (4.7 in), with females growing larger than males.

Habitat and Biology

This species is very common along the eastern U.S. coast, from the Gulf of St. Lawrence to northern Florida. Mummichogs are especially abundant in salt marshes and tidal creeks of the ACE Basin and coastal South Carolina. The spawning season extends from spring to fall and varies with latitude. During this time, male mummichogs become brighter in color. Spawning takes place during the new or full moon (when the tide is at its highest) in water of various salinities. Eggs may be laid in a variety of substrates, depending on availability. Mummichog eggs may be found in the empty shells of ribbed mussels, on the leaves of marsh grass, in pits dug out and covered by the female, or spread directly on the bottom. The eggs develop out of the water, and hatching takes place upon immersion during the successive moon tide. Mummichogs are extremely hardy and can tolerate widely fluctuating environmental conditions. During winter months, they may burrow in the mud or move to deeper water at the mouths of channels. The word "mummichog" comes from an Indian word meaning "going in crowds." Various age classes will gather, often close to shore, in schools that may number several hundred fish. Feeding occurs at the marsh surface, in mid-water, or on the bottom even though this species' upward tilted mouth is specialized for surface feeding. Mummichogs eat a variety of foods including marine worms, crustaceans, small shrimps, insects, and other fish.

Species Significance

Mummichogs are an important food source for many commercially valuable fish species, as well as wading birds and seabirds. They are an important link in the coastal food chain. Mummichogs have little commercial (small industry for selling as bait) and no recreational value. Mummichogs and other species of killifish are used extensively in scientific studies, including medical and environmental research, and have been introduced to many areas for controlling mosquito larvae.

References

Abraham, B. 1985. Species profiles: Life histories and environmental requirements of coastal fishes and invertebrates (mid-Atlantic)--Mummichog and Striped Killifish. U. S. Fish and Wildlife Service Biological Report 82 (11.40). U. S. Department of Interior, Washington, D.C.

Boschung, H.T,. Jr., J.D. Williams, D.W. Gotshall, D.K. Caldwell, and M.C. Caldwell. 1983. The Audubon Society field guide to North American fishes, whales and dolphins. Alfred A. Knopf, New York, NY.

Murdy, E.O., R.S. Birdsong, and J.A. Musick. 1997. Fishes of the Chesapeake Bay. Smithsonian Institution Press, Washington, DC.



Red Drum Sciaenops ocellatus

red drum

SoundSound, SCDNR

Description

The red drum, also called "channel bass" or "spottail bass," is a member of the drum family, Sciaenidae, and a close relative of Atlantic croaker, spot, and kingfish. Red drum have elongate bodies, reddish-bronze color, and a distinct spot or spots at the base of the caudal fin. Scientists believe that the prominent spot on the tail confuses predators into attacking the fish's tail instead of its head, thus giving the fish a better chance to escape. Similar to other members of the drum family, the red drum has an inferior mouth and a lateral line that extends to the tip of the caudal fin. Also, red drum males produce "drumming" sounds during the spawning season to attract females. Males also undergo a color change during this time, becoming dark red or bluish gray on top with a pale underside.

Habitat and Biology

Along the Atlantic coast of the United States, red drum are found from Delaware to southern Florida. They also inhabit the Gulf of Mexico, from southern Florida to northeastern Mexico. In South Carolina, red drum can be found in estuarine areas, including those in the ACE Basin. Red drum utilize different habitats as they develop. The large, adult fishes do not live in the same habitat as the subadults. In general, adults are found in nearshore and coastal waters, whereas subadults are usually found in larger creeks and rivers, although they have been observed in waters off barrier islands and sandbars. Juveniles are abundant in the shallow creeks that meander through cordgrass (Spartina alterniflora) marshes. As juveniles mature, their habitat preferences change and they move from the shallows of the estuaries into deeper inlets and along the front beaches. Because of difficulty in sampling the adult stock, very little is known of their biology. After spawning i n August and September, the adults can be found around the front beaches and inlets. As water temperatures decline in the fall (late October-November), the adults move gradually offshore to deeper, warmer water. This offshore movement initiates a flurry of successful surf fishing. The movement is repeated in the opposite direction in the spring. As inshore water temperature warms in the spring, the red drum move from the deeper, warmer offshore waters to the inlets and front beaches, where they are once again available to anglers.

In general, individuals found inside the state's estuaries are juveniles; that is, they have not as yet reached sexual maturity and are frequently called subadults. Males mature at age three (68.6-76.2 cm or 27-30 inches long); females mature at age four (81-91 cm or 32-36 inches long). Reproductive activity in this species coincides with cooling temperatures and shorter days in late August. Spawning in South Carolina waters is thought to occur in coastal inlets, including St. Helena Sound, although some nearshore spawning activity is believed to occur in South Carolina during August and September. At this time, male red drum produce characteristic "drumming" sounds by contracting muscles near the swim bladder, in an effort to attract females. Fertilized eggs are pelagic, and hatching occurs in 28 to 29 hours, depending on water temperature (longer in cool water and shorter in warmer water).

red drum larvae Upon hatching, larvae face the difficult task of reaching nursery grounds in the upper reaches of estuaries. They are transported to the mouths of estuaries in currents generated by wind and tides. By swimming towards the surface when currents are rising (flood tide ) and staying close to the bottom when the tide is ebbing so as not to be swept back out to sea, they eventually reach the shallow tidal creeks, where they settle out of the plankton community and spend the first few months of life. As larvae, after yolk sac reserves are exhausted, red drum feed on zooplankton in the water column. Once in the estuary, larval red drum feed on crustaceans and small fishes. The smallest fish feed on copepods, and as they grow, they eat ghost shrimp known as mysids and eventually consume grass and penaeid shrimps. As waters cool in December, these small fishes (2.54-7.62 cm or 1-3 inches long) leave the shallow marsh creeks for the relative warmth of deeper creeks and estuarine rivers. When water temperatures rise in the spring, small red drum re-enter these tidal marsh creeks and begin to grow very rapidly. At this time they feed on small fishes and crustaceans. In May and June, these 20.3-25.4 cm (8-10 inch) fish, along with fish of previous year classes, begin to leave the small creeks to inhabit the open estuarine shallows. At one year of age, red drum are 25.4-30.48 cm (10-12 inches) long, and weigh just under .45 kg (1 pound). Most meet the minimum legal size of 35.56 cm (14 inches) in October (13 months of age) and weigh on average slightly more than .45 kg. By December (15 months of age), many of these fishes may weigh almost .9 kg (2 lbs).

Many of the 1 to 3 year old red drum inside the estuary show an interesting pattern of movements and feeding that is related to the stage of the tide. During the warmer months of the year, as the incoming tide begins to reach the smooth cordgrass, fish move into the grass, where they feed on fiddler crabs (80% of their diet), mud crabs, grass shrimp, and fishes that are associated with this structured habitat. As the tide ebbs, the red drum move off the marsh surface and are generally found on mud bars with oyster reefs in shallow water adjacent to the marsh. In the winter when water temperatures are low and the fiddler crabs are no longer active, red drum display the same pattern of tidal movement, but they become sluggish and less active since they are unable to regulate their body temperature. During the warmer months, their movements reduce their exposure to bottlenose dolphin, which is a major predator. In the coldest part of the year, they tend to be more vulnerable to predation by this marine mammal.

Species Significance

In South Carolina, red drum are classified as a gamefish and can only be bought or sold through a mariculture operation or imported from a state that allows commercial fishing. Moreover, this species can only be harvested by hook and line and by gig during the appropriate season. Drum that are less than 14 inches and are caught in South Carolina State waters must be released by law. Also, there is a bag limit of five fish per angler per day, only one of which can be longer than 27 inches. (See related section: Recreational Fisheries.)

In recent years, the red drum has been placed among the nation's important recreational and commercial species which have been fished into a state of decline. Red drum numbers in South Carolina have increased in recent years, but commercial fishing is still restricted. The fisheries for red drum have been conducted almost entirely within the internal waters of the states; therefore, management has traditionally been by individual state regulation. In South Carolina, restrictions have been issued to help manage red drum populations.

References

Mercer, L.P. 1984. A Biological and fisheries profile of the red drum, Sciaenops Ocellatus. Special Scientific Report No. 41. North Carolina Department of Natural Resources and Community Development, Division of Marine Fisheries, Morehead City, NC.

Vaughan, D.S. and T.E. Helser. 1990. Status of the red drum stock of the Atlantic coast: Stock assessment report for 1989. National Oceanic and Atmospheric Administration, Beaufort, SC.

Wenner, C.A. 1992. Red drum: Natural history and fishing techniques in South Carolina. Educational Report No. 17. South Carolina Department of Natural Resources, Marine Resources Division. Charleston, SC.

Wenner, C.A., W.A. Roumillat, J.E. Moran Jr., M.B. Maddox, L.B. Daniel III, and J.W. Smith. 1990. Investigations on the life history and population dynamics of marine recreational fishes in South Carolina. Part I. Report to Fish Restoration Act under Project F-37. South Carolina Department of Natural Resources, Marine Resources Division. Charleston, SC. .



Redbreast Sunfish Lepomis auritus

Redbreast Sunfish

Description

Redbreast sunfish are close relatives of well-known fish species such as the large- and small mouth basses, bluegill, and crappie, all of which are members of the family Centrarchidae. Sunfish typically have compressed bodies and a spinous dorsal fin followed by a soft-rayed fin. These two dorsal fins are joined, so they appear as one. Identification of sunfish is often difficult, especially when closely related species mate to produce hybrids. Adult redbreast sunfish are brownish above with olive green or violet hues. Their heads have lines and spots of a pale blue color, and the flap of the gill cover is black with a pale yellow edge. The sides of the fish are greenish with vertical bars that tend to be more distinct in females than in males. The underside is yellowish white, except during breeding season, when it becomes bright orange in the males and yellow in the females (hence the redbreast sunfish's common name). Redbreast sunfish are more elongated than other Lepomis species and generally attain lengths of up to 25 cm (10 in). The current South Carolina record is 0.9 kg (2 lbs).

Habitat and Biology

This species is commonly found in freshwater habitats from Maine to Florida and westward along the Gulf Coast to Texas. Sunfish are common in the upper blackwater reaches of all rivers and streams in the ACE Basin. The sunfish can survive in a variety of environmental conditions, such as headwater streams to coastal plain rivers and lakes. They prefer flowing water and are most often found associated with stumps or logs in mainstream habitats, but can exist in impoundments. They can survive in lakes and streams at elevations of up to 1,000 meters (3,500 feet) and waters less than 8 ppt. Redbreast sunfish can withstand pH ranges from 4.8 to 8.4; however, they cannot survive in waters warmer than 37° C.

Spawning generally occurs in May and June in the ACE Basin. The full moon of May is the "traditional" time of the peak spawning season. Spawning occurs over sandy or gravel bottoms in lakes, ponds, streams and rivers. When spawning in rivers, this species tends to do so in faster flowing water than other sunfishes. The nest consists of a circular depression on the bottom that is lined with pebbles. Most often it is associated with some type of cover and is in the main channel of the stream. Occasionally, redbreast sunfish will occupy the nests of other species. They do not have community nests like other Lepomis species. Males guard the eggs until hatching and are continually fanning the nest to increase oxygen levels and remove siltation, Sexual maturity is achieved in the second year, and females can lay up to 14,000 eggs. Most redbreast live to an age of three or four years.

Similar to most sunfish, redbreast sunfish are sight feeders and capture food either by lying in wait and making a sudden lunge or by actively foraging along the bottom. Preferred food items of redbreast sunfish include aquatic insects, small crustaceans, and fish. They most often feed in the middle of the water column. They are often associated with a particular haunt, such as a submerged tree, rock or overhanging bank. The abundance of snag habitats that provide a constant source of aquatic insects in a system is an important factor in regulating population health.

Species Significance

Redbreast sunfish are an important recreational species in South Carolina. According to creel census conducted in the Edisto and Combahee Rivers during 1987-91 by the South Carolina Department of Natural Resources, redbreast sunfish were the most popular fish among anglers during late spring and early summer, dominating harvests in terms of abundance. They are a forage fish for other predators such as largemouth bass. Predation by flathead catfish may have significant impacts on sunfish populations.

References

Allen, D.E. and C.S. Thomason. 1993. Survey and inventory of fisheries resources in the Combahee River. Study Completion Report F-32. SC Wildlife and Marine Resources Department, Columbia, SC.

Carlaner, C.D. 1977. Handbok of freshwater fishery biology. Vol. 2. Iowa State University, Ames, IA.

Fisheries Habitat Committee. 1995. Fishes of the Edisto River basin. SC Department of Natural Resources, Water Resources Division Report 6. Columbia, SC.

Jenkins, R.E. and N.M. Burkhead. 1994. Freshwater fishes of Virginia. American Fisheries Society, Bethesda, MD.



Sheepshead Archosargus probatocephalus

Sheepshead

Description

Sheepshead have stout, oval-shaped bodies with short heads, the most obvious feature of which are rows of broad, peg-like teeth (hence their common name). They are grayish in color with five or six broad black, vertical bars on either side of the body and an incomplete head bar. These bars are usually more distinct in younger animals. The dorsal fin is continuous, with sharp, strong spines preceded by a small spine that points forward and is embedded in the skin. The second spine on the anal fin is large and very strong. The pectoral fins are greenish and pointed and extend past the anal opening when pressed against the body of the fish.

Habitat and Biology

Sheepshead spawn in near-coastal waters during the winter and early spring (December - March). During this time they are frequently seen on artificial reefs, as well as natural reef habitats in depths of 18 to 22.5 m (60 to 75 feet). This species is a member of a fish family (Sparidae) more frequently found in offshore waters. Members of the Sparidae commonly are hermaphroditic and undergo sex-reversal. Sheepshead, however, apparently show no evidence of hermaphroditism; each mature individual functions as a male or female throughout its entire adult life.

Eggs and larvae are transported to the estuaries, and small sheepshead are recruited to the shallow tidal creeks that meander through the Spartina marshes. They prefer the higher salinity creeks that have an abundance of oysters and submerged structure. The young sheepshead are cryptic; that is, they are attracted to structure and use it as a hiding place. Little is known about their early life history, but young sheepshead can frequently be caught near dock pilings or other submerged structures. Submerged structures are covered with a community of encrusting organisms such as barnacles, bryozoans, etc. The pattern of dentition of sheepshead enables them to effectively pick or scrape encrusting organisms from underwater structures from the time they are small juveniles through their adult lives. Throughout life, they feed on attached fauna such as barnacles and mussels, as well as small crabs and shrimp.

Sheepshead display a seasonal migration in the fall from inshore higher salinity waters to offshore waters. The pattern is reversed in the spring when, as the inshore waters warm, they move back into the higher salinity parts of the ACE basin. They are relatively long-lived, with the oldest individual reported having an estimated age of 23 years. They grow quickly during the first three years of life; but after reaching sexual maturity, the growth slows dramatically.

Species Significance

Sheepshead are sold commercially in South Carolina, although no directed commercial fishery exists. Commercial landings result from incidental catch. They are, however, an important recreational species in South Carolina waters. Anglers who target sheepshead seek them near structures such as pilings or oyster bars. This species is not currently threatened or endangered, and there are no size or bag limit restrictions on it.

References

Bearden, C.M. 1961. Common marine fishes of South Carolina. Bears Bluff Lab Contribution No. 34. Bears Bluff Laboratory, Inc., Wadmalaw Island, South Carolina.

Boschung, H.T,. Jr., J.D. Williams, D.W. Gotshall, D.K. Caldwell, and M.C. Caldwell. 1983. The Audubon Society field guide to North American fishes, whales and dolphins. Alfred A. Knopf, New York, NY.

Johnson, G.D. 1978. Development of fishes of the mid-Atlantic Bight: an atlas of egg, larval and juvenile stages. Volume IV: Carrangidae through Ephippidae. U.S. Fish and Wildlife Service, Office of Biological Services FWS/OBS-78/12. Ft. Collins, CO.

Roumillat, W. 1998. pers. comm. South Carolina Department of Natural Resources, Charleston, SC.



Shortnose Sturgeon Acipenser brevirostrum

Sturgeon

Description

Sturgeons are fish of an ancient lineage easily recognized by five rows of scutes (bony plates) along their bodies: one row along the mid-back, one along the middle, and one along the lower body on each side. Sturgeons have heterocercal tails; that is, the top lobe of the caudal fin is larger than the bottom one. Coloration varies from yellowish pink to yellowish brown on the fish's back and creamy white below. Sturgeons, the largest of the bony fishes, are bottom dwellers that use chin barbels to locate food on the substrate. The barbels on the shortnose sturgeon are rather small, less than one half the width of the mouth. Sturgeons have protrusible, inferior mouths used to suck in benthic insects, crustaceans, and other food items. The shortnose sturgeon is smaller than the common Atlantic sturgeon, Acipenser oxyrinchus, and has a shorter, uncurved snout. It is also known as the blunt-nosed, round-nosed and small sturgeon, and it may grow up to 143 cm (56 in).

Habitat and Biology

Shortnose sturgeons are found from the Saint John River, Canada (New Brunswick), to the St. Johns River, Florida. They inhabit estuarine and riverine habitats and are typically not found offshore. In South Carolina they inhabit Winyah Bay Rivers, those that drain into Lake Marion, the Santee, Cooper and Savannah rivers, and the ACE Basin. In the latter, shortnose sturgeon are typically found at the freshwater-saltwater interface. Adult and sub-adult shortnose sturgeon are known to inhabit this area during spring through fall. Spawning may take place well upriver; however, the existence of a spawning stock in the ACE Basin is yet to be determined.

Shortnose sturgeons are semi-anadromous; they migrate from the estuary into rivers to spawn. Adults migrate upstream in early spring and forage in the lower reaches of rivers at the fresh tidal water and estuarine water interface. This area is also an important nursery habitat for juveniles, which do not usually migrate. Migration of shortnose sturgeons and the extent to which they utilize freshwater habitats vary throughout the species' range. For instance, in northern latitudes, sturgeon make use of freshwater habitats more extensively than in southern regions, and some do not frequent estuarine habitats at all throughout the year. Sturgeons can live more than 50 years and typically grow very slowly. Growth and age at sexual maturity vary with latitude. Fish from southern locations grow faster and mature at a younger age than fish in the north. Male sturgeons inhabiting South Carolina waters become mature at approximately age 4 and females at age 6-7. Male sturgeons repro duce 1-2 years after reaching maturity. However, females may not spawn until 5 years after becoming mature. In South Carolina, the age at which female sturgeons first spawn ranges from 7 to 14 years. Spawning periodicity varies among individuals, but spawning rarely takes place in consecutive years. A female typically produces 27,000 to 208,000 eggs and can carry an egg mass of over 1 million eggs, depending on her body size. Spawning in South Carolina occurs from February to April over gravel or rubble bottoms. High current velocity and adequate substrate for the attachment of eggs are important factors in spawning site selection. Sturgeons are benthic feeders preferring mollusks and crustaceans as adults and insects and small crustaceans as juveniles.

A related species, the Atlantic sturgeon, is anadromous, spawning in freshwater rivers and spending the rest of its life in estuarine and nearshore waters. Females may spawn only every 2-6 years. Spawning in South Carolina begins in February and continues through the spring, with a second spawning migration possibly occurring in the fall. Spawning populations of Atlantic sturgeon in South Carolina are found in the Savannah River, one or more of the rivers in the ACE Basin, the Santee River, Winyah Bay and its river systems, and possibly the Cooper River. Although definitive spawning locations have not been identified, spawning is believed to occur in tidal freshwater, in deep areas with high water flow rates. A "marl hole" approximately 15 meters (50 ft) deep above Givhans Ferry on the Edisto River may be one of these locations. Females may release between 400,000 and 3.7 million eggs during a spawning period. As the eggs are released, they are fertilized externally by a male and then sink to the bottom. Sturgeon eggs are sticky and will adhere to rocks, plants, and other solid material. The eggs hatch within a week or so. Within three weeks, juvenile sturgeon have absorbed their yolk sac and begun to feed on bottom-dwelling organisms such as small shrimp, worms, clams, snails, and small demersal fish. Juvenile Atlantic sturgeon spend their first few years in rivers and estuaries with some movement along the coast. Tagging studies in South Carolina indicate that higher growth rates occur during the winter, spring and fall, with slower growth rates during the summer. As the juvenile sturgeon reach 0.6 to 0.9 meters (2-3 ft) in length (age of 1-6 years) , they begin to move into oceanic waters and join the adult population, remaining there until sexual maturity (7-19 years and nearly 2 meters or 6-7 ft in length). Atlantic sturgeon may undergo long migrations up and down the coast.

Species Significance

Worldwide, sturgeons are commercially valuable as a source of high-grade caviar, and their meat is popular both smoked and fresh. In the past, both Atlantic and shortnose sturgeons were reported in the landings for sturgeon. In the United States, Atlantic sturgeon supported a commercial fishery; however, landings have not exceeded 300,000 lbs. since 1920, and all Atlantic states have closed their fisheries in recent years. The sturgeon fishery was closed in South Carolina in 1985 following precipitous declines in numbers landed in the early 1980s. Shortnose sturgeons are currently of no commercial value because of their status as an endangered species. There is no recreational fishery for this species in the United States.

Shortnose sturgeon were listed as endangered throughout their range under the Endangered Species Conservation Act of 1969. In 1967, the U. S. Fish and Wildlife Service cited pollution and overharvesting as primary factors for the decline in numbers of shortnose sturgeon. Other sources contributing to population declines include incidental catch in shad gillnet fisheries, dam and bridge construction, dredging, entrapment in power plant water intake screens, and reservoir operations. In South Carolina, the primary factors affecting populations of this species are habitat alteration, due to dredging and dam construction, and pollution. Construction of dams has the potential to reduce suitable spawning sites, and disturbance associated with dredging activities impacts the food supply for juvenile sturgeon.

References

Collins, M.R. and T.I.J. Smith. 1993. Characteristics of the adult segment of the Savannah River population of shortnose sturgeon. Proceedings of the Annual Conference of the Southeastern Association of Fish and Wildlife Agencies 47:485-491.

Dadswell, M.J., B.D. Taubert, T.S. Squires, D. Marchette, and J. Buckley. 1984. Synopsis of biological data on shortnose sturgeon, Acipenser brevirostrum. Technical Report 14. National Oceanic and Atmospheric Administration, National Marine Fisheries Service, Seattle, WA.

Hall, J.W., T.I.J. Smith, S.D. Lamprecht. 1991. Movements and habitats of shortnose sturgeon, Acipenser brevirostrum, in the Savannah river. Copeia 1991(3):695-702.

Leland, J.G., III. 1968. A survey of the sturgeon fishery of South Carolina. Bears Bluff Laboratory Contribution No. 47. Bears Bluff Laboratory, Wadmalaw, SC.

Murphy, S. H. and G. Barnette (eds.). South Carolina's endangered species portfolio. South Carolina Department of Natural Resources, Wildlife Diversity Section, Columbia, SC.

Shortnose sturgeon recovery team. 1997. Recovery plan for the shortnose sturgeon-public review draft. Office of Protected Resources, National Marine Fisheries Service. National Oceanic and Atmospheric Administration, Silver Spring, MD.

Taub, S.H. 1990. Fishery management plan for Atlantic sturgeon. Fisheries Management Report 17. Atlantic States Marine Fisheries Commission, Washington, DC.



Silver Perch Bairdiella chrysoura

Silver perch

Description

Silver perch are among the smaller members of the drum family (Sciaenidae) and common in South Carolina estuaries. They have oval-shaped bodies and no barbels. Their coloration is relatively plain, with an olive, green or bluish back and a silvery white belly. Fins are all yellowish, with the dorsal, caudal and sometimes the tip of the anal fin somewhat darker. Silver perch have a few strong spines at the angle of the preopercle, the largest of which point down.

Habitat and Biology

Silver perch also belong to the sciaenid family. They spawn during the late spring and early summer inside the estuary, usually in the deeper sections of smaller tidal creeks in the higher salinity areas. Eggs and larvae remain in the estuary, and when larvae become capable of active swimming, they settle in the shallow tidal creeks. This habitat is also the silver perch's nursery area, and the young feed on small decapod crustaceans and fishes. During the fall, silver perch move from the shallow marsh habitat into the deeper creeks and main part of the estuaries of the ACE Basin. During high tide, the larger silver perch move into the tidal creeks, where they feed on grass shrimps. As the tide drops, they move back out into deeper water.

Species Significance

No commercial or recreational fisheries exist for this species. However, pier fishers often catch silver perch and use them for bait or for personal consumption. They are a forage fish for larger predatory fish such as spotted seatrout and red drum and also are an important food source for many birds.

References

Bearden, C.M. 1961. Common marine fishes of South Carolina. Bears Bluff Lab Contribution No. 34. Bears Bluff Laboratory, Inc., Wadmalaw Island, South Carolina.

Johnson, G.D. 1978. Development of fishes of the mid-Atlantic Bight: an atlas of egg, larval and juvenile stages. Volume IV: Carrangidae through Ephippidae. U.S. Fish and Wildlife Service, Office of Biological Services FWS/OBS-78/12. Ft. Collins, CO.

Murdy, E.O., R.S. Birdsong, and J.A. Musick. 1997. Fishes of the Chesapeake Bay. Smithsonian Institution Press, Washington, DC.

Wenner, C. 1998. Personal communication. South Carolina Department of Natural Resources, Charleston, SC.



Southern Flounder Paralichthys lethostigma

Southern Flounder

Description

The southern flounder is one of approximately 130 American flatfish species. Southern flounder belong to the family Paralichthyidae. Flatfishes are compressed laterally (flat), with the dorsal side of the body pigmented in order to match their surroundings. Southern flounder are light to dark brown with darker spots and blotches that are not well defined. All flatfishes have eyes that migrate dorsally during larval development, so both eyes are found on the "ocular" side. The ocular side in southern flounder is the left side, but in other flatfishes it may be the right side. The underside of the fish, the ventral or "blind" side, is usually light in color. Maximum adult size in this species is 75 cm (30 in).

Habitat and Biology

Southern flounder are a benthic species found in coastal waters of the southeastern United States, from the Chesapeake Bay to Florida (except southern Florida), and along the Gulf of Mexico. They can tolerate a wide salinity range and commonly inhabit brackish and freshwater habitats. Females grow faster than males; each sex matures as they approach their second birthday (males are 20.3-25.4 cm or 8-10 inches long; females are 30.5-35.6 cm or 12-14 inches long). The average life span of a male is two years, and the oldest male southern flounder reported in South Carolina was age 3+. Females are dramatically larger than males at a given age and a three-year old female may weigh 0.7 to 2.7 kg (1.5 to 6 pounds).

flounder larvaeBeginning in late September of each year, southern flounders that are older than one year begin to mature and leave the estuaries for offshore spawning grounds. The locations of these offshore spawning grounds are unknown at this time. Males leave the estuaries and move towards the spawning grounds earlier than females. Their migration coincides with a 4°C to 5°C drop in temperature. Only the young-of-year that are approaching age one remain inside South Carolina estuaries from December through April. After they spawn and water temperatures rise, mature fish re-enter the estuaries. After hatching, the larvae feed and grow near the ocean surface as they are transported toward shore by currents initially associated with the Gulf Stream. The swimming ability of the juveniles, coupled with the presence of tidal currents in near-shore waters, provides a mechanism for the transport of the young to the estuarine nursery habitat. Settling (affiliation with creek bottoms and attainment of adult pigmentation ) of the southern flounder occurs in lower and moderate salinity areas of South Carolina estuaries from January through March of each year. This species utilizes the same shallow marsh habitat as the red drum and spotted seatrout as a nursery area. In the shallow tidal creeks they feed on small shrimps and fishes. As they increase in size, fish become the preferred prey. Southern flounder remain in these creeks until they attain a length of about 20.3-25.4 cm (8-10 inches), at which time they disperse into other available estuarine habitats. They can even tolerate fresh water and have been taken by electrofishing.

Southern flounder are ambush predators with the ability to change coloration so that they blend in with the substrate. They are frequently found in shallow water near oyster reefs or the mouths of small rivulets draining the high marsh as the tide ebbs. Food items include grass shrimp, mummichogs, spot, and striped and white mullet. These prey animals are commonly associated with edge habitats in the estuary such as the edges of oyster bars and along salt marshes. Southern flounder are widely distributed in the ACE Basin and frequently co-occur with red drum.

Species Significance

Southern flounder are popular in both the commercial and sport industries in the Southeast and Gulf of Mexico. In the latter, much of the commercial catch of southern flounder results from bycatch from the shrimp fishery. Recreationally, southern flounder are caught by nighttime gigging in tidal creeks and marshes, as well as by hook and line

References

Fernandez, E. 1991. The juvenile life history of the southern flounder (Paralichthys lethostigma) in South Carolina. M.S. Thesis. University of Charleston, Charleston, SC.

Murdy, E.O., R.S. Birdsong, and J.A. Musick. 1997. Fishes of the Chesapeake Bay. Smithsonian Institution Press, Washington, DC.

Reagan, R.E., Jr. 1985. Species profiles: Life histories and environmental requirements of coastal fishes and invertebrates (Gulf of Mexico)--Southern Flounder. U.S. Department of Interior, Fish and Wildlife Service, Biological Report 82(11.30). U.S. Army Corps of Engineers, Report TR EL-82-4. Washington, DC.

Wenner, C. 1998. Personal communication. South Carolina Department of Natural Resources, Charleston, SC.



Southern Kingfish Menticirrhus americanus

whiting

Description

The southern kingfish, or whiting, is a member of the family Sciaenidae and a common inhabitant of South Carolina estuaries, including those in the ACE Basin. It has an elongated body and a characteristic single barbel on its chin. The mouth is small and inferior, as is characteristic of fishes that feed on the bottom. Coloration varies somewhat with habitat, but it is generally gray to brown above with silvery reflections and 6-8 broad oblique lateral bars. The latter are not always distinct, however. Fins are dusky, sometimes with a black margin, especially on dorsal and pectoral fins. The caudal fin of adult kingfish has an elongated lower lobe, making its margin appear "s" shaped. Pelvic, anal and caudal fins sometimes exhibit a yellowish coloration.

Habitat and Biology

This species spawns in the late spring and summer in near-shore coastal waters. Young juveniles occur in channels and along the coast in the penaeid shrimp fishing grounds. Males and females are sexually mature by age two, and the oldest individual reported in South Carolina is age five. Their absence in the coldest months of the year suggests that they move either south or slightly offshore to warmer waters. Southern kingfish feed on a variety of benthic infauna and epifauna such as polychaetes, crustaceans and mollusks.

Species Significance

In South Carolina waters, there is no directed commercial fishery for southern kingfish; the commercial catch is incidental to the penaeid shrimp-trawl fishery. Though not considered a gamefish by recreational anglers, whiting are an excellent food fish and are sought after by pier fishers and surf fishers throughout the state. This species is neither threatened nor endangered, and there are presently no size or bag limit restrictions on it.

References

Bearden, C.M. 1961. Common marine fishes of South Carolina. Bears Bluff Lab Contribution No. 34. Bears Bluff Laboratory, Inc., Wadmalaw Island, South Carolina.

Johnson, G.D. 1978. Development of fishes of the mid-Atlantic Bight: an atlas of egg, larval and juvenile stages. Volume IV: Carrangidae through Ephippidae. U.S. Fish and Wildlife Service, Office of Biological Services FWS/OBS-78/12. Ft. Collins, CO.

Smith, J.W. and C.A. Wenner. 1985. Biology of the southern kingfish in the South Atlantic Bight. Transactions of the American Fisheries Society 114:356-366.



Spot Leiostomus xanthurus

Spot

Description

Spot, which belong to the drum family (Sciaenidae), derive their common name from the presence of a dark spot or blotch behind the upper end of the opercle or gill slit. They have a short, flattened body with an elevated back, and, unlike Atlantic croaker, they lack barbels. Spot are purplish gray above, with a golden luster and oblique streaks that extend below the lateral line. Their caudal fin is darkish, while the other fins are usually pale yellow. In early fall, as spawning season approaches, South Carolinians often refer to the large males as "golden spot." Presumably, this is because males tend to acquire a more defined golden hue during this time of the year.

Habitat and Biology

Spot are found along the east coast from Florida to the Middle Atlantic states during the warmer months. These fishes are estuarine dependent, with a variety of habitats being utilized by various life history stages in the estuaries of the ACE Basin. In fall, spot migrate south and offshore to spawn during late fall and early winter in the warmer waters caused by the presence of the Gulf Stream. Larval spot use ocean and tidal currents to transport them from the offshore spawning areas to the estuarine nursery habitat. As they approach the coastal inlets, spot rise to the surface during the flood tides and move through the water column towards the bottom during ebb tide. By using differential current velocities from surface to bottom (surface currents are stronger than those near the bottom), spot gain access to the estuaries.

Although numerous authors have described the estuary as a nursery for small fishes and shellfish, not all habitats within the estuary are used by a given species. The shallow marsh habitat, with its series of dendritic creeks, mudflats, and shallows, is the principal nursery area for spot after they gain access to the estuary. They recruit to these shallows via tidal currents and settle from the water column to become demersal. In general, the smallest spot are found further upriver and in shallower water. The headwater regions of salt marshes are not only a refuge from predators but also have an abundant food supply for juveniles. Spot apparently live in association with estuaries and shallow coastal waters until they attain maturity (about 2 years old), at which time they migrate to deep ocean waters to spawn. Spot feed on benthic infauna, primarily polychaetes, crustaceans, and mollusks.

Species Significance

Spot are harvested commercially throughout their range. Along the east coast, the commercial catch of spot has averaged 3.7 million kg (8.2 million lbs) per year since 1950. Spot are also a large constituent of the by-catch from the South Atlantic penaeid shrimp-trawl fishery. Recreational catches throughout the region averaged 1.7 million kg (3.9 million lbs) per year since 1981. In South Carolina waters, recreational anglers target spot mainly in the fall, as the fish move out of the estuaries towards offshore spawning grounds. Spot are abundant and ubiquitous inhabitants of South Carolina waters. They are a regular part of the diet of many of the predators in the estuary such as spotted seatrout, red drum, herons, and osprey. There are no size restrictions or bag limits governing the harvest of spot in South Carolina.

References

Bearden, C.M. 1961. Common marine fishes of South Carolina. Bears Bluff Lab Contribution No. 34. Bears Bluff Laboratory, Inc., Wadmalaw Island, South Carolina.

Johnson, G.D. 1978. Development of fishes of the mid-Atlantic Bight: an atlas of egg, larval and juvenile stages. Volume IV: Carrangidae through Ephippidae. U.S. Fish and Wildlife Service, Office of Biological Services FWS/OBS-78/12. Ft. Collins, CO.

McErlean, A.J., S.G. O'Connor, J.A. Milhursky, and C.I. Gibson. 1973. Abundance, diversity and seasonal patterns of estuarine fish populations. Estuarine Coastal Marine Science 1:19-36.

Pacheco, A.L. 1962. Age and growth of spot in Lower Chesapeake Bay, with notes on distribution and abundance of juveniles in the York River system. Chesapeake Science 3(1):18-27.

Potter, I.C., P.N. Claridge, and R. M. Warwick. 1986. Consistency of seasonal changes in the estuarine fish assemblage. Marine Ecology Progress Series 32(2-3):217-228.

Shenker, J.M., and J.M. Dean. 1979. The utilization of an intertidal salt marsh creek by larval and juvenile fishes: Abundance, diversity, and temporal variation. Estuaries 2(3):154-163.

Weinstein, M.P. 1979. Shallow marsh habitats as primary nurseries for fishes and shellfish, Cape Fear River, North Carolina. Fisheries Bulletin U.S. 7:339-357.

Wenner, C. 1998. Personal communication. South Carolina Department of Natural Resources, Charleston, SC.



Spotted Seatrout Cynoscion nebulosus

spotted seatrout

Description

The spotted seatrout is not a member of the trout family (Salmonidae), but of the drum family (Sciaenidae) which includes common inhabitants of South Carolina waters such as Atlantic croaker, red drum, and spot. Two other species of seatrouts inhabit state waters: weakfish, Cynoscion regalis, and sand seatrout, Cynoscion nothus. All seatrouts are elongate with a protruding lower jaw. Their mouths are rather large with sharp teeth. Spotted trout are gray with bluish reflections above and silvery below. The fins are yellowish green, except for the first dorsal fin which is dusky. They can be distinguished from other seatrouts by numerous black spots on their upper side, second dorsal fin, and caudal fin. Maximum adult size in this species is 90 cm (35 in).

Habitat and Biology

Spotted seatrout are found along the eastern coast of the United States from Chesapeake Bay to the southern tip of Florida and along the Gulf coast to the northeast Mexican coast. Spotted seatrout are estuarine dependent, and with the exception of forays into the nearshore waters during the warmer months of the year, they spend their whole lives inside estuaries. Male spotted seatrout mature at a much smaller size than females. Regardless of the month in which a male spotted seatrout was spawned the previous year, it will become mature and actively spawn the following May (as small as 19 cm or 7.5 inches). Female spotted seatrout do not become mature until they attain a size of about 25.4 cm (10 in) in length, approximately one year after spawning. Females spawned near the end of the spawning season of the previous year will not mature until the following spawning season.

 trout larvae Seatrout spawn at specific locations in South Carolina's estuaries from May through early September. Typically, these sites have swiftly moving currents with obstructions on the bottom and are either in high salinity areas or are near the inlets. During the spawning season, mature males produce a "drumming" sound by contracting muscles near the swim bladder. All fish belonging to this family produce these noises (hence the common name of "drums"). Females are attracted by the drumming to areas where large numbers of males are gathered. Spawning occurs in such gatherings, or aggregations, near dusk. Larvae are transported by tidal currents to the shallow estuarine habitat such as dendritic tidal creeks, muds flats, oyster bars, and intertidal marshes. In these tidal creeks, juvenile spotted seatrout feed on crustaceans, mainly grass shrimp, and small fishes. They remain in this habitat until reaching a length of about 6 inches after ,which they move to larger, deeper estuarine creeks. In the coldest months of the year (December through early March), young-of-year spotted seatrout move to the deeper holes in the smaller creeks and on the edges of the main channels. As the waters warm in the spring, they form schools of like-sized individuals and move throughout the estuary feeding on small fishes and shrimp.

Spotted seatrout approximately 33 cm (13 in) feed on small fishes and shrimps found in shallow waters near flooded marsh grass, and as they grow, fish become the preferred prey. Males grow more slowly than females and reach a smaller maximum size. The oldest spotted seatrout aged in South Carolina was 8 years old.

Species Significance

The spotted seatrout is one of the most popular sport fish along the eastern coast of the United States and Gulf of Mexico. It ranks second by weight among recreational saltwater anglers mainly in the southeastern United States. In South Carolina, the species was officially declared a gamefish on July 1, 1986. Trout can only be harvested by means of hook and line or by gig in South Carolina waters. Hook and line harvest is allowed throughout the year, whereas use of gigs is permitted only from March through November. Currently, there is a size limit of 33 cm (13 in) and a bag limit of 10 fish per angler per day.

The Marine Division of the South Carolina Department of Natural Resources, in conjunction with the National Marine Fisheries Service, recently reported that the total recreational catch for spotted seatrout averaged 266,000 individuals per year over an 11-year period. The population of spotted seatrout in state waters is declining, albeit slowly, due to increased fishing pressure. However, changes in the way the spotted seatrout resource is managed in South Carolina and other Atlantic states can ensure a stable future for seatrout populations.

References

Boschung, H.T,. Jr., J.D. Williams, D.W. Gotshall, D.K. Caldwell, and M.C. Caldwell. 1983. The Audubon Society field guide to North American fishes, whales and dolphins. Alfred A. Knopf, New York, NY.

Murdy, E.O., R.S. Birdsong, and J.A. Musick. 1997. Fishes of the Chesapeake Bay. Smithsonian Institution Press, Washington, DC.

Wenner, C. 1998. Personal communication. South Carolina Department of Natural Resources, Charleston, SC.

Wenner, C.A. and J. Archambault. 1996. Spotted seatrout: Natural history and fishing techniques in South Carolina. South Carolina Department of Natural Resources, Marine Resources Research Institute Educational Report 18.



Striped Bass Morone saxatilis

striped bass

Description

Striped bass are members of the family Percichthyidae, the temperate basses. They are elongated with 7-8 dark stripes extending horizontally, a dark olive to steel blue back and silver underside with a brassy sheen. The two dorsal fins are separated by a gap, and two spines are present on the edge of the opercle. The caudal fin, or tail, of striped bass is clearly forked. Males reach a maximum length of 116 cm (45 in), whereas females grow to about 183 cm (72 in). Maximum weight is about 57 kg (125 lbs) with the South Carolina record being over 23 kg (50 lbs). In 1965, South Carolina biologists were able to cross, or hybridize, white bass (Morone chrysops) with striped bass. The resulting fry survived, and since then, other hybrids have been produced between striped bass and other species of the genus Morone. White bass are smaller than both striped bass and its hybrid. They are light green to gray above and silvery below with no distinct lines or stripes and have a small head and arched back. Striped bass are the largest of the three. As described above, their coloration is dark, they have a large head and mouth, and the back is not arched. The hybrid looks like a combination of the two, with the characteristic back-arch of the white bass, a small head, and dark gray or blue body and stripes that are frequently broken.

Habitat and Biology

Striped bass are native to the Atlantic coast, from the St. Lawrence River, Canada, to the St. John's River, Florida. On the Gulf coast, it is distributed from the Suwannee River, Florida, to eastern Texas. Because striped bass can live in fresh water for long periods of time, they are stocked in many inland reservoirs. However, only two East Coast reservoirs have self-sustaining populations: the Kerr Reservoir in Virginia and North Carolina, and the Santee-Cooper Reservoir in South Carolina.

Striped bass are native to the ACE Basin. They belong to the southern strain and behave quite differently from their northern relatives. Southern fish, unlike northern fish, never leave their riverine environments. Northern fish spend a considerable amount of time in near-shore waters and then ascend the rivers to spawn. Striped bass in the ACE Basin never enter the ocean, and it is strongly suspected that they never leave the river in which they are born. Striped bass are found in all the large rivers of the ACE Basin, and they over-winter in the estuarine areas of these systems near the saltwater-freshwater interface. Summers are spent in the cooler waters of the upper river, where springs and a dense canopy of trees keep water temperatures lower. They are often found in deep holes in the river or around structures such as old pilings.

Striped bass are schooling, anadromous fish; that is, they spend much of their time in salt water but migrate to fresh water rivers to spawn. However, landlocked populations spend their entire life in freshwater and do not migrate. Bass from North Carolina and the Chesapeake Bay are known to undertake coastwide migrations in addition to annual spawning migrations. They move north to New England and Canada during early spring and return between September and December. Bass inhabiting waters south of Cape Hatteras, North Carolina, typically do not take part in coastal migrations. Recent advances in molecular genetics have allowed researchers to investigate differences in populations of striped bass. Evidence strongly indicates that the rivers of the ACE Basin contain a population of striped bass that is unique to the basin.

In South Carolina, spawning migrations typically begin in March, when water temperatures exceed 58°F, and continue through early summer, with males arriving at spawning grounds before females. Fish move to areas just upstream of the saltwater-freshwater interface, where the female releases her eggs to be fertilized by any pursuing males. The semi-buoyant eggs then drift in the tidal currents for several days until they hatch. Spawning success is often sporadic because of the limited range of environmental conditions required for eggs to hatch and larvae to grow. Sexual maturity occurs around the fifth year at about 71 cm (28 in) in length. Eggs are pelagic, and larvae hatch in approximately 2-3 days. Larvae depend on endogenous nutrition for the first 5-10 days. Endogenous nutrition means that larvae derive nutrients and energy from the material contained in their yolk sacs. After this stage, once larvae have well-developed mouths, they begin to feed on zooplankton . Juveniles feed on a variety of worms, small crustaceans, insects, and fishes. In freshwater impoundments, fish such as herring, alewife, and shads constitute the main diet of the adult striped bass, while fishes, squid, clams, lobsters, crabs, shrimps, and other invertebrates are taken by adults in open-water habitats.

Species Significance

This species of bass has historically been America's most important recreational and commercial fish. Sportfishing attracts many fishermen to South Carolina. Historically, the commercial industry for striped bass has added millions to the state's economy, being an important income and employment opportunity in the coastal areas. Hatcheries exist throughout the East Coast. This bass species is also used as a "biological control" to regulate gizzard shad and herring populations in large reservoirs.

In the 1950s, the striped bass population in South Carolina exploded, causing an increase in recreational fishing, especially in the Santee-Cooper Reservoir. This population growth eventually caused a decrease in striped bass's feeding fish, the herring and shad, causing their populations to plummet. Consequently, the striped bass population in the Santee-Cooper also began to decline due to starvation. This "boom-bust" cycle is common to many fish species. A significant decline of striped bass throughout the East Coast began in the late 1970s. The U.S. Congress responded to this in 1979 by amending the Anadromous Fish Conservation Act to include an emergency striped bass study. In 1981, the ASMFC published an Interstate Fisheries Management Plan for Striped Bass. In 1980 the Striped Bass Study was implemented to identify possible causes of the decline of striped bass and to outline an action plan and research program to address these causes. Possible causes of fish decline include over-harvesting, habitat deterioration, contaminants, and industrial development. In 1984 the U.S. Congress passed the Atlantic Striped Bass Conservation Act, requiring a federal moratorium on striped bass fishing in those states which have not adopted the recommended management measures of the ASFMC Plan or are not satisfactorily enforcing these measures.

References

Deuel, D., D. McDaniel, S. Taub. 1989. Atlantic coastal striped bass: Road to recovery. U.S. Department of the Interior, Fish and Wildlife Service, and U.S. Department of Commerce, National Oceanic and Atmospheric Administration. Washington, D.C.

Fay, C.W., R.J. Neves, and G.B. Pardue. 1983. Species profiles: Life histories and environmental requirements of coastal fishes and invertebrates (Mid-Atlantic)--Striped Bass. U.S. Department of Interior, Fish and Wildlife Service, Biological Report FWS/OBS-82/11.8. U.S. Army Corps of Engineers Report TR EL-82-4. Washington, DC.

Fuller, J.C., Jr. 1984. South Carolina's striped bass story. South Carolina Wildlife and Marine Resources Department, Columbia, SC.

Hardy, J.D., Jr. 1978. Development of fishes of the mid-Atlantic Bight: an atlas of egg, larval, and juvenile stages. Volume 3: Aphredoderidae through Rachycentridae. U.S. Fish and Wildlife Service, Office of Biological Programs. FWS/OBS-78/12. Ft. Collins, CO.

Jenkins, R.E. and N.M. Burkhead. 1994. Freshwater fishes of Virginia. American Fisheries Society, Bethesda, MD.

Williams, H.M. 1975. Characteristics for distinguishing white bass, striped bass and their hybrid (striped bass X white bass). South Carolina Wildlife and Marine Resources Department, Abbeville, SC.



Striped Mullet Mugil cephalus

striped mullet

Description

Mullet (family Mugilidae) are common inhabitants of freshwater, estuarine and offshore habitats and are well known for their habit of jumping, sometimes up to 1 meter (3 feet), out of the water. Striped mullet, also called grey mullet and black mullet, are elongate with a series of dark stripes that run the length of the body. The dorsal, or upper part of the fish's body, is a bluish-gray color, the underside is white, and there is a purplish blotch on the upper base of the pectoral fins. Similar to other mullets, striped mullet possess a well-developed adipose eyelid, a thick membrane that covers part of the eyeball and presumably functions in protection. Striped mullet have two well-separated dorsal fins (the first one with four spines), pectoral fins that join the fish's body above the midline (towards its back), and no lateral line. The mouth, when viewed from below, appears as an inverted "V."

Habitat and Biology

This species of mullet is found in temperate and tropical waters throughout the world. Along the western Atlantic, striped mullet are distributed from Brazil to Nova Scotia, including the Caribbean and the Gulf of Mexico. Adult mullet aggregate in large schools and migrate from shallow estuarine areas and freshwater rivers to offshore spawning grounds in the fall and winter. The spawning season for this species is from October to February, with peak activity in November-December. After hatching, larvae remain offshore until they reach about 20 mm (< 1 inch) in length. At this time, they begin migrating back to shallow coastal areas, such as estuaries, where they complete their development. In the fall, young mullet either move to deeper waters or remain in estuaries to spend the winter. Once they become adults, mullet are found in various habitats including freshwater rivers, saltmarshes, estuaries, and the open sea. The feeding habits of striped mullet vary with age. Young mullet feed primarily on small crustaceans and other zooplankton, whereas adults ingest plant matter. They feed by sucking in bottom sediments that contain decaying plant material, algae, and inorganic particles. They may also extract algae and microorganisms from scum that accumulates on the water surface or from the surface of submerged vegetation or other substrates. Swarming marine worms (polychaetes ) have also been reported as part of their diet.

Species Significance

Striped mullet have both commercial and recreational value. They are commonly used as bait for favored sport fishes, such as billfish, but they are also a popular food fish in the southeastern United Sates and Gulf coast, where they are sought after for both their meat and their roe (egg mass). The heaviest commercial fishing for mullet is done during spawning migrations, when fish are gathered in large schools near the surface. Gill nets and cast nets are used in the commercial harvest of mullet, whereas recreational fishermen find cast netting to be particularly effective since mullet tend to congregate near the surface.

Striped mullet are neither threatened nor endangered. However, in recent years, the Japanese demand for mullet roe has increased, causing American stocks of this species to decline sharply.

References

Bearden, C.M. 1961. Common marine fishes of South Carolina. Bears Bluff Lab Contribution No. 34. Bears Bluff Laboratory, Inc., Wadmalaw Island, South Carolina.

Boschung, H.T,. Jr., J.D. Williams, D.W. Gotshall, D.K. Caldwell, and M.C. Caldwell. 1983. The Audubon Society field guide to North American fishes, whales and dolphins. Alfred A. Knopf, New York, NY.

Collins, M.R. 1985. Species profiles: Life histories and environmental requirements of coastal fishes and invertebrates (South Florida)--striped mullet. Biological Report 82(11.34). U.S. Fish and Wildlife Service, Washington, DC.

Murdy, E.O., R.S. Birdsong, and J.A. Musick. 1997. Fishes of the Chesapeake Bay. Smithsonian Institution Press, Washington, DC.

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