The Life History of Red DrumGary C. Matlock

Texas Parks and Wildlife Dcpartrnent4200 Smith School Road

Austin, Texas 78744

Thc red drum Sciaenops ocellatus Linnaeus! is aquasi-.catadromous sciaenid Rounsefell, 1975! thatranges from Tuxpan, Mexico in the Gulf of Mexico Gulo to Massachu set ts in the Atlantic Ocean Hild<»brand and Schroeder, 1928; Bigelow and Schroedcr,1954; Simmons and Breuer, 1962!. I mporta n t d irectedfisherics for red drum have existed since the 1700s,but are mainly limited today to the Carolinean prov-ince Briggs, 1974; Matlock, 1980!. In Texas, red druidare harvested primarily in estuaries where they oncecomprised as much as 35 percent of the commercialfish landings. Rccrea tiona1 fishermen may reta in fivered drum per day in Texas.

Major conflicts have occurred over allocating reddrum for 100 years Ma tlock 1980!. Recreational fish-errnen blame inadequate catches on commercialoverfishing, and commercial fishermen accuse sportfishermen of having inadequate skill. Managementusually attempted to reduce conflict by addressingrecreational concerns without appreciably affectingcommercial harvests Matlock, 1980!. Prior to the1970s, the success of each regulation was ~arely ex-amined, and procedures for selecting them wereseldom specified. Size limits were enacted in the1920s to protect juvenile and adult fish Pearson,1929!, and seasonal and area net closures werc ex-panded over time to protect spawning adults Hef-fernan and Kemp, 1980!.

In Texas, addihonal regula tions resulted until thesale of native red drum was prohibited in September1981 because of overfishing Maflock, 1982!. Man-agement's goal was inodified from reducing conflictto obtaining optimum yield OY! Anonymous, 1982!,

Identity and Morphology

Eggs and Yolk-sac LarvaeOnly laboratory-spawned specimens have txvn

described Johnson et al., '1977; Holt et al., 'I981a,1981b!, but both eggs and yolk-sac larvae are similarto those of other sciaenids in that they are small and

From "The Basis for the Development of a lVfanagement Planfor Red Drum in Texas," the author's Ph.D. dissertation forTexas A&M University. College Station, Texas.

lack pigmt:ntation, which makes fiel identificationunreliablebclow the Family level Marlcy,1983!. Thespherical eggs are 0.86-0,98 mm in dianicter, andusually contain onc colorless oil droplc t about 0.2.-0,4 mm in diameter Johnson ct al. 1977!, have aperivitelline space less than 2 percent of the cggdiameter, and have a clear, unsculpturcd chorion,

I.a rva c are 1,71.-1,79 mm standard length a t hatch-ing. [Illustrations of red drum eggs and larvae arepresented in Growth and Development of Red DrumEggs and Lathe elsewhere in this book. j

Larvae

Rcd d ru m larvae arc difficult to identify becausethey are morphologically similar to Atlantic croaker Micropogonias undulatus! except for pigmentationand anal fin spines. Sciaenids considered to be onespecies based on descriptions af Pearson �929! andHildebrand and Schroedcr �928! werc caught in theTexas surf, reared in the laboratory, and later identi-fied as both Atlantic croaker and red dru m Siinmonsand Breuer, 1962!. However, recent descriptions oflaboratory-spawned red drum larvae Johnson et al.,1977! are consistent with previous pigmentationdescriptions Pearson, 1929; Lippson and Moran,1974; Hildebrand and Cable, 1934!; niyomcre andana! fin ray descriptions Pcarson, 1929; Miller andJorgcnsen, 1973!; illustration of a sharp opercularspine Jannke, 1971!; and illustration of caudal finclcments Topp and Cole, 1968!. Fin developmentand pigrnenta tion of laboratory-reared fi sh Johnsonet ul., 1977! were not consistent with the description ofa similar sized individual given by Powlcs and Sten-der �978!. Descriptions of laboratory-reared fish areprobably thc most reliable because they are based onmany fish from known parents.

Juveniles, Sub-Adults and AdultsFxternal morphology and swim bladder and oto-

lith shapes identify juveniles, sub-adults and adults.The young have two short tube-like diverticula onthe carrot-shaped swim bladder, characteristics ofthc supragencric group Sciaenops Chao, 1978!. Thesediverticula remain as fish mature and a pair of sac-like projections develop dorsolaterally in the anteriorswiin bladder. These projections fit into a body wal!

Red L>rum Aquaculture

cavity between the third and fourth pleural ribs andmay be involved in sound reception in older fish. Theotolith's sagitta has the sciacnid characteristics of atadpole-shaped sulcus in its inner surface but is en-larged and slightly rectangular. Chao �978! summa->~zcd thc external morphology as: snout with fiveupper and five marginal pores; lower jaw with fivepores; no barbel or lower jaw; mouth inferior; teethvilliform in bands; and gill rakers short.

The juveniles and adults differ externally in cau-dal fin shape and cxtcrnal color. Caudal fins arepointed in young and become slightly concave inadults Pearson, 1929!, Large black blotchcs are dis-tributed over the sides and back in fish 100 mm I fildcbrand and Schroeder, 1928!. A pronouncedchromatophore enlargement at the upper caudal finbase appears at about 36 mm and remains through-out life. Lateral blotches enlarge until the fish reachabout 150 mm, then fade and disappear, Adults areelongate, silvery reddish fish with an elevated back Hildebrand and Schroeder, 1928; Pearson, 1929!.Thc head is long, rather low, with a bluntish snoutand a larger inferior mouth; body shape may be anadaptation to shallow surf habitat Chao, 1978!.

Reproductive CycleSpawning

Courtship behaviorRed drum arc dioccious, and males court fcrnalcs

before spawning, Drumming and nudging intensi-I'ics prior to spawning and may bc a major stimulusfor spawning, at least in the laboratory Guest andLasswcll, 1978!.

Spa tvni ng periodicitySpawning occurs from August through January

and peaks in September or October. This conclusionseems supported by both field and laboratory evi-dence, much of which is not well documented. Sexu-ally mature adults, larvae, and small fish <150 mm!occur mainly in fall Table I!. Moreover, only simu-lated fall photoperiod and temperatures inducedspa wning in the labora tory Arnold et al., I 977; Robertsetal.,1978a!. Thedateofpeakspawningvariesamongyears, but bag seine collections in bays indicate Octo-ber I as a reasonable hatching date for modelingpurposes Ma tlock, 1984!.Spau ning areas

No study has comprehensively examined spawn-ing areas, but probable areas can be suggested by aprocess of elimination using the criterion that theabsence of rcd drum from larval collections probablyindicates no spawning at the collection sites.

Spawning apparently docs not occur in bays orthe offshore gulf. Larvae do not occur beyond 20 km�8 m depths! from gulf beaches Houde et al., 1979;Arnold et a!., 1976; Finuca neet al., 1977!. They are also

absent from Aransas, Corpus Christi, and AlazanBays H<>ese, I 965; Holland et al., 1973, 1974; Dokken,1981! where grass beds, channels, shorelines, areasnear scrpu lid rock formations, and deep water wereextensively, alth >ugh not randomly, sampled, InMississippi, larval abundance increased as distancefrom bays increased out to 20 km Laroche, perse>nalcomment!, On the Atlantic coast, bay and offshorecollections also lack larval red drum Pearson 194 1;Massmann ct al. 1961, 1962; Herman 1963; Crocker1965; Dovcl 1967!, although most of these collectionswere made north of Virginia in areas where few reddrum now occur.

Spawning seems to occur in the nearshore gulf,probably near bay-gulf passes, Many larvae �.11mm! have been captured in bay-gulf passes Comp-ton, 1965; Jan nkc 1971; King, 1971 !. In contrast, onlyone larval red drum was caught in two years ofichthyoplankton sampling in the Gulf �8 to 135 mdeep! off Texas; that larva was caught at the shallow-est �8 rn! station Arnold et al., 1976; Finucane et al.,1977!. These recent studies confirm the nearshorc

gulf spawning site originally proposed by Pcarson�929! and supported by Simmons and Breuer �962!and Yokel �966! because they caught spent adultsalong gulf beaches and felt juveniles werc mostabundant near passes. Their cvidcnce for this wasweak, however, because: I ! the occurrenceof ripe fishin bays was ignor>4, as were adult movements beforeand after spawning; 2! juvenile catches werc notadjusted for distribu tion of sampling effort, althoughmostcollcctions werc ncarbay-gulf passes; 3! spawn-ing was not observed, and eggs were not collected;and 4! the offshore gulf was not sampled. Although"ripe" fish ha ve been reported in bays Pearson, 1929;Yokel, 1966!, this tnay not indicate where spawningoccurs because fish like red drum cruise at aboutthree times their length per second Harden Jones,1968!. So, a 750-mm adult could travel 194 km/d.Moreover, the term "ripe" was not defined and mayor may not indicate imminent spawning.

Hydrographic considerations also suggest gulfspawning, probably easterly or northerly of estuar-ies. The larvae reported from passes are probablytransported by nearshore gulf surface currents whichare strongly along shore to thc south and southwestand have an onshore component during fall. In addi-tion, spawning outside estuarine nurseries is consis-tent with the triangular migration pattern typicallyexhibited by marine fish in which larvae are trans-ported from spawning areas to spatially separatenurseries, and adults occupy separate feeding oroverwintcring grounds Harden Jones, 1968!.

Recruitment and Nurseries

Egg and larval transportGulf surface currents probably carry eggs and

Af at la ck � Li fe History

larvae from ncarshorc spawning areas to estuarinenurscrics. Evidence to support this includes: I! nddrum eggs fioa t in gulf salinities �5 to 35%ho! Holt etal., 1981b!, 2! nearshore surface currents in the gulfduring fall are onshore and strongly along shore in asouth to southwest direction Watson and Behrens,1970; Armstrong et al., 1976!; and 3! larvae concen-trate at the surface in Texas passes King, 1971!. Thesame conclusion was reached for the sand scatrout,Cynoscionarenarius, Shlossman and Chittendcn,1981!another estuarine sciaenid. Moreover, Standard andChi t tcndcn �984! and Murphy and Chit tcndcn �982!have su g gested tha t spawning in non-estuarine I'ishessuch as the banded drum, primus fasciatus, and theGulf butterfish, Peprilus burti, is also correlated withsouth/southwest along-shore current transport atthat time of year and that it may reflect a generalcorrelation between spawning periodicity and cur-rent transport. Howcvcr,gulfhydrologyisnotundcr-stood to the extent that existing physical oceano-graphic models can predict exactly where larvaecaught in bay-gulf passes were spawned. Brucks,personal communication!.

Subsurface currents may also be involved in lar-val transport within passes and bays, but supportingevidence is weak and the relationship may dependon the type of estuary. Mansueti �960! suggestedthat distribution within Chesapeake Bay, a two-layerfiow with vertical mixing type estuary Bowden,1967!, was determined by subsurface currents, al-though he did not have adequate current data. Alongthe gulf, where most estuaries are the verticallyhomogeneous type, vertical distribution of larvae isinconsistent. Jannke �971! reported greatest catchesOf larvaC near bOttOm in Florid, but King �971!found greatest catches at the surface off Texas. Rea-sons for this apparent difference in thc gulf are notclear, although wind conditions may strongly influ-ence mixing in vertically homogeneous estuaries.

Nursery locationsNo study has comprehensively examined nurs-

ery locations, but probable nurseries can be sug-gested by a process of elimination using the criterionthat thc absence of juveniles and sub-adults from alocation probably indicates that area is not a nursery.

Red drum do not usc the offshore gulf as a nurs-ery. Dcspitc extensive collections in depths from 4 to400 m, juvenile and sub-adult fish have rarely, if ever,been captured with: 1! shrimp trawls Hildebrand,1954; Springer and Bullis, 1956; Miller, 1965; Chitten-den and McEachran, 1976; Chittenden and Moore,1977; Bryan et al., 1982; Ross et al., 1982!; 2! fish tra wls Cody et al., 1977!; 3! midwater trawls C.E. Bryan,personal communication!; 4! longlincs Cody et al.,1981; G. Graham, Texas A&M University unpub-lished data!; 5! rods and reels C.E. Bryan, personalcommunication; McEachron and Matlock, 1983!; 6!

fish traps C.F. Bryan, personal communication!; or7! purse scincs Knapp, 1950; Christmas et al., 1960;Dunham, 1972!.

Juveniles and sub-adults only occasionally occurin thc surf zone. Chittcndcn personal communica-tion! captured I 0 fish about 300 mm on hook and linewithin 8 m of shore off Big Shell Beach on Padreisland in August1980, a week after Hurricane Allen,This appears unusual, however, and may be relatedto tha t hurrica nebecause more extensive seine collec-tions rarely yield juveniles in the surf Gunter, 1958;McFarland, 1963; Juneau, 1975; Moddc and Ross,1981!.

Rcd drum utilize estuarine nurseries. Many sci-entists have captured large numbers of juveniles andsub-adults in estuaries Mansucti, 1960; Yokel, 1966;Jannkc, 1971; Matlock et al�1978; Hegen, 1982; Mat-lock, 1983; Holt et al,, 1983!. A tolerance to low salini-ties that increases with larval age Crocker et al., 1981!may be an adaptation to estuarine conditions.

Distribution and Movements

JuvenilesJuvenile rcd drum occur in all Gulf and Atlantic

estuaries from Chesapeake Bay to thc Laguna Madreof Mexico. However, their abundance among Gulfestuaries varies, depending mainly on surface area Yokel, 1966!. Salinity may also affect distribution,probably by affecting survival in early life stages Neff et al., 1981; Wohlschlag, 1981!, Although rainand river in flow cause salinities to bc lower along theupper coast of Texas �7+/-4 %a annually in GalvestonBay! than on the lower coast �5 +/- 6 %~ in upperLaguna Madre! Matlock, 1984!, juvenile abundanceis similar.

Bay-gulf passes interconnect estuarine nurscricswith Gulf spawning grounds and areas occupied byadults. Each estuary in Texas, except theupper LagunaMadre and San Antonio Bay, is connected to the Gulfby at least one pass Brown et al., 1974!. These twobays are connected to the Gulf indirectly by the Intra-coastal Waterway. Additional connections may becreated when hurricanes crcatc ncw passes or tempo-rarily re-open old ones.

Little information exists on distribution of juve-niles within estuaries, They apparently prefer shore-lines, shallow water <1,3 m!, and sea-grass mead-ows. Collections in opcn-bay areas seldom yieldjuveniles Mansueti, 1960; Jannke, 1971!, but shore-line collections typically do Hegen, 1982!. Althoughreasons for their distribution are not know~ becausethe literature essentially describes only their pres-ence and absence, red drum occur throughout thebays, over all substrates, in vegetated and nonvege-tated areas, adjacent to human dcvcIopmcnts, inmarshes, and in channels and rivers during winter Simmons and Brcuer, 1962; Yokel, 1966; Pcrret et al.,

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1980!. Holt et al. �983! found fish �- 27rnm SL! preferthc area between sea grass Halodu!e sp.! and non-vcgctatcd bot toms.

Sub-adults

Sub-adijlts occur in the Gulf only occasionally,Recreational fishermencatch sub-adults in the surf in

thc fa Il but only infrequently Weixelman, 1982!. Fcwstudies report sub-adults from the surf zone or thcGulf scc Nursery Areas section for references!,However, McFarland �963! seined three 440-mmfish in thc Gulf during the day in summer 1960 andspeculated morc might be caught at night.

Sub-adults apparently rema in in est ua riesthroughout the year and do not migrate seasonallybetween estuaries and gulr. Many tagged fish havebeen recaptured in winter in estuaries Osburn et al.,1982; Creen et al., 1985!. In addition, fishery-inde-pendent gill and trarnrnel net catch rates Gunter,1945; Matlock et al., 1978; Hegen and Matlock, 1980;Hcgen, 1981!, recreational catch rates Heffernan etal., 1977; Brcucr et al., 1977!, and commercial land ings Hamilton, 1980, 1981, 1982! either remain thc samethroughout the year or generally decline after recruit-mcnt of new year classes in fall.

The distances that sub-adults move are small, and

these fish normally remain within one estuary. Fisharc usually recaptured within 10 km of the taggingsite Table 2; Carr and Chancy, 1976; Adkins et al.,1979; Osburn et al. 1982!, so red drum do not exhibitthc broad random movement that Moe �972! re-ported for sciaenids in general. More than 77 percentof 1,339 tag returns were from the estuary wheretagging occurred Osbu m et al., 1982!; 95 percent ofall recaptures included fish that had only moved toadjacent estuaries.

Little information exists on distribution of sub-

adults within estuaries, although temperature-in-duced movements do occur between shallow anddeep water. Like the juveniles, sub-adults are distrib-uted throughout each estuary. However, their habi-tat preferences are unknown. Vegetation and reeftypes and amounts vary among bays, but relativeabundance generally does not Matlock, 1984!. Sub-adults are more abundant along shorelines or shal-low water <L3 m! than in open bay in fall and spring,but will temporarily move to water >L3 m to escapetemperature extremes in winter Matlock et al., 1978;Matlock, 1979!. Matlock et al, �978! reported highestgill net catches adjacent to deep holes just prior topassage of cold fronts indicating these fronts movefish to deep water within estuaries. Pearson �929!,Gunter �945!, and Miles 951! reported red drumemigrate from estuaries to the Gulf in winter, but thiswas based only on absence of fish in shoreline net setsthen and appears wrong based on Matlock et al.�978!. Gunter's �945! trammel net catches, rnore-over, were equally high in winter �.4 fish/set! and

summer �.1 fish/sct!. Saline gradients may helporient localized movements required for behavioralthermorcgulation Owens et al., 1981!.

ln the laboratory sub-adults arc most active atnight, Owens et at �981! reached this conclusion bycontinuous electronicmonitoringof movements,butthe validity of extrapolation to wild fish is unknown.

Adults

Red drum permanently emigrate from estuariesto the gulf when sexually mature, Sexual maturity isprobably reached in 3,5 years Matlock,1986!, Adults >750 mrn! rarely occur in the estuaries Matlock,1984!, but commonly occur in the Gulf Wcixclman,1982; Ross et al., 1982!. This presence in the Gulfseems permanent because they are absent from bays.Moreover, Simmons and Breuer �982! found that allof 30 returns from 134 fish �00-460 mm! tagged atCedar Bayou, Texas, in April were from thc Gulf.

Adult red drum in the Gulf are distributed to atleast 113 km from shore. Most occur within 16 km ofshore off Texas,but current literature is inadequate todescribe in detail the distribution of adults in the

Gulf, Charter boa ts catch these fish off Louisiana nearoil rigs to 34 km offshore in 13 m deep water Dugaset al., 1979!, and on reefs 113 km off Texas TexasParks and Wildlife Department, unpublished data!.Fish were caught in purse seines about 19-21 km offthe Chandeleur Islands, La., in 18 m deep water Na tional Marine Fisheries Service, unpublished da ta!and with fish trawls, bottom longlines, and shrimptra wls with in 24 km of f Texa s Cody et al,, 1977; Cod yand Avent, 1980; Cod y et al,, 1981; Ross et al,, 1982!,

Feeding Behavior artd DietFeeding

Red drum feed throughout the water column, butmainly at the bottom. This conclusion is based on thefollowing: 1! they eat mostly bottom organisms in thewild; 2! fish 279-312 mm FL fork length! eat live foodinaquaria only when it ison a substrate Yokel,1966!;and 3! fish searching for food in tanks apparentlyorient their body to the bottom by contacting it withtheir lower jaw and small pelvic fin ray extensions Yokel, 1966!. Prey are sucked up from the bottom bya rapid expansion o f the branchial region or capturedby biting the substrate.

Red drum feeding in shallow water often exhibit"tailing" behavior. ln this activity their caudal anddorsal fins are out of the water as they feed on thebottom Gunter, 1945; Simmons and Breuer, 1962;

Yokel, 1966!, Schools of tailing fish can be easilysighted which increases their vulnerability to bayfishermen.

Occasionally rcd drum feed at the surface, Schoolshave been sighted close to Gulf beaches feeding onother fish at thc surface Rohr, 1968!. I have seen

M<itlo< k � l ife History

schools of rcd druin feedirig at the surface on Gulfmenhaden Brevovrtia patroni<s! at hvated water dis-charges in Trinity Bay, Fish demonstrating this be-havior are also easily sighted which probably in-creases their susceptibility to fishcrincn,

Diet

Rcd drum feed indiscriminantly within at leastthree ontogenetic stages and, after yolk-sac absorp-tion, their diet is va ried. Overstrevt and Heard �978!found 59 taxa in juveniles, sub-adults and adultsfiom Mississippi Sound. The most common taxawcie crustaceans and fishvs, as found in many otherstudies Table 3!, Boothby and Avault �971! consid-ered red drum to bc omnivorous feeders because 69percent of 197 stomachs examined contained morethan two foods. Even marsh rats have been reported Pcarson, 1929!. As with most sciaenids, prey availa-bility and size probably determines diet Chao andMusick, 1977; Boothby and Avault, 1971!.

Larvae begin exogenous feeding within four daysof spawning and eat mainly zooplankton. The yolk-sac provides nourishment for the first four days post-spawn Johnson et al., 1977!. Bass and Avault �975!determined preference by simultaneously samplingzooplankton and fish and found fish �0 mm prefercalanoid and cyclopoid copelx>ds while those 30-70mrn prefer mysids, Copepods are the common foodof fish <25 mrn regardless of geographic location Colura and Hysrn 1th, 1976; Od um, 1971!. Ho wever,red drum 25-50 rnm will utilize what is available inthe absence of mysids Colura and Hysrnith, 1976!.

Juveniles mainly cat small bottom invertebratesand young fish. I'ish 60-100 mm rely heavily onarnphipods Bass and Avault, 1975!. Pcnaeid shrimp,callinectid crabs, and fish menhaden, gobies, mullet,killifish, and eels! prcdorninate in fish larger than 100mm Table 3!.

Adults eat mainly fish. Fish occur more frequentlyin comparison to invertebrates as red drum grow Pcarson, 1929; Yokel, 1966; Overstreet and Heard,1978! .

Male and female diets seem similar. Boothby andAvault �971! reported no noticeable dict differencebetween sexes, but presented no supporting data.Rcd drum dict apparently changes among seasonsand geographic locations. However, the change maybe caus<xi more by prey availability or size than byseason. For example, shrimp and fish replaced crabsin winter in similarly sized �90-780 mrn! MississippiSound fish Table 3! when crabs were less abundant Overstrcet and Heard, 1978!. Similar change in taxaoccurred in summer at Hopcdalc, La. Table 3!. Asfish grew through sumrncr in earthern ponds poly-chactcs and amphipods replaced copcpod s Table 3!.In Carninada Pass, La,, dict changed dramatically Table 3! within seasons as prey availability and sizechanged Bass and Avault, 1975!.

Rvd drum dictmaydifferbetwcen day and nightdepending on fish size, Fish 35-40 mm ate mainlymysid s in both day and night Bass and Avault, 1975!.However, fish 90-115 mm ate mainly palaemonidshrinip in the day and fish at night.

ln summary, rcd drum diet varivs with their size,temporally, and spatially probably bn ause of preyavailability and size of prey change,

Parasites and Diseases

The literature on red drum parasitvs and diseasesdeals primarily with their identification. [In SectionIV of this manual, S,K, Johnson describes the lifehistory and control of several parasites and diseasescommon to rcd drum.] Thirty-one organisms havebeen found on or in red drum, but copcpods andccstodcs arc most frequent Table 4>. The occurrcnccof many species may be related to the omnivorousfeeding habits of rcd drum Yokel, 1966!.

Infections have generally not been quantified,Yokel �966! concluded that rcd drum are not heavilyparasitize%,but the studies he summarized generallyfailed to give the number of fish examined or infectedor thc number of parasites found.

Estimates of Life History ParametersFecund i ty

Females probably produce more than 500,000eggs annually. Existing estimates are 551,600,-3,420,552 eggs/female Table 5!, However, they arebased on few fish, undescribed techniqu es, or do notconsider fish size.

Growth of Juveniles and Sub-AdultsMuch information exists on juvenile and sub-

adult growth, Thcirgrowth hasbccn determined by;1! measuring length changes in! abora tory and pond-rcared fish Colura et al., 1976; Arnold et al., 1977;Johnson et al�1977; Roberts et al., 1978b; Trirnble,1979; Hcin and Shepard, 1980; Crockvr et al., 1981;Holt et al., 198l a; Hysmith et al., 1983! and in taggedwild fish Siinmons and Brcucr, 1962; Matlock andWeaver, I 979b; Perret et al. 1980; McKcx., 1980!; 2!leng th-frequency analysis of'wild fish Pea rson, 1929;Miles 1951; Simmons and Brcucr, 1962; Bass andAvault, 1975!; 3! counting rings annuli! on scales ofwild fish Welsh and Breder, 1924; Pearson, 1929;Wakefield and Colura, 1983!, and 4! counting ringson otoliths of wild fish Thieling and Loyacana, 1976;Rohr, 1980!. However, growth is still not accuratelyknown because the application of laboratory andpond studies to the natural environment may not bevalid, and most studies on wild fish Table 6! areunreliable because they arc based on few fish, inade-quate dcscriphon of procedures, inadequate statisti-caI analysis, gear selection bias, questionable data,and failure to adjust for time of annulus formation.

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Although much growth information exists, fcwmathematical expression~ relating length and age ofred drum have been developed. Rohr �980! Wakc-field and olura �983! estimated parameters in thcvon Bertalanfiy cquatiori using otoliths and scalesfrom Louisiana and Texas. However, Rohr publishedonly an abstract, and Wakeficld and Colura collectedfish from a few bays only.

Juveniles in Texas reach 90-95 mm at 6 mor>thsand I'15-150 mm at 8 months Matlock, 1984!. Juve-niles ger>eral!y grew in size continuously and in anexponential manner during their first 300 days with-ou t slowing during winter, Sub-adults mach 330-360inm at 18 months, and 550-600 mm at 26 months Matlock,1984!. Thescgrowthestirnatcs agrccclosclywith that based on the von Bertalanffy equation forTexas fish based on Pcarson's 1929 data! but not forSouth Carolina fish Matlock, 1984!. Values for K,L, and t were 0.295, 1068 mm, and + 0.144/ycar Texas! and 0.449, 945 mm, and -0.324/year SouthCarolina!. Predicted length at age for the von Berta-lanffy equations were 106, 352, and 480 mm Texas!and 292, 528, and 636 inm South Carolina! at 6, 18,and 26 months,

Seemingly exponential growth explained morethan 68 percent of the variation in juvcnilc lengths Matlock, 1984!. Their growth may not really bcexponential, however, because gear selection overes-timates length carly and underestimates it late ineach phase. The estimate that size at 0.8/year is 160-240 mm Ma tlock, 1984! may be too low, bu t it agrees~ ith the von Bertalanffy length estimate of 190 m mat0.8/year using Pearson's data. Mean daily growthduring the first 8 months of life �.53-0.80 mm! issimilar to published rates Table 6! that approximate0.6 mm/d in the wild and 1.0 rnm/d in controlledlaboratory, raceway, or pond environment wheregrowth may be higher.

Reported values of K in the von Berta la nffy equa-tion are 0.3 Pcarson's data!, 0.4 Rohr 1980!, 0.35-0.52 Wakcfield and Colura 1983!, and 0.45 Thicling andLovacano's data!. The von Bertalanffy cquationbasedon Pearson's data may be most reasonable bccausc:I! L from it is below but reasonably close to thelargest fish on record in Texas; 2! length at ages oneand two years estimated trom it arc consistent withlengths from bag seines and traminel net collections;3! his da ta were based on the greatest size range; and4! he included fish collected in the gulf where thelargest fish occur. Wakefield and Colura's �983! andThicling and Loycano's �976! estimates were basedon data from estuaries and may reflect size biasbecause old large fish emigrate to the Gulf whereasRohr's estimate may reflect unreliable otolith read-ings in old fish, These authors estimate L as beingless than 950 mm which may reflect inadequate rep-resentationn of old fish, because many fish larger than950 mm occur in the Gulf.

MortalityMuch work has been doneon red drum mortality.

Howcv«r, except for recent work on total mortalityby !vlatlock and Weaver �979b! and Greeri et al.�985!, parameters needed for yield assessments havenot been emphasized. Vast work has emphasizedcauses of infrequent massive kills Gunter, 1941, 1952;Guntcr,ind Hildcbrand, '1951; Simmons, 1957; Sim-mons and Hreiier, 1962; Bryan, 1971; Harrington,1973; Cardcilhac et al., 1981; Amos, 1981!, influence oftemperature and salinity on hatching succc'ss andlarval mortality Holt et al., 1981a; Crocker et al.,1981!, larval cannibalism Arnold et al,, 1977!, effectsof the [XTOC!oil wellblowouton larvalmorphologyand survival Rabalais et al., 1981!, and parasites anddiseases of age onc year and older fish Yokel, 1966!The availablc literature on red drum predators pro-vides little insight into mortality because few preda-tors have been identified Gunter, 1942, 1945; Kemp,1949; Reid et al., 1956; Darnell, 1958; Caldwell andCaldwell, 1972; Diener ef a!., 1974; Moffett ef al., 1979;C>vcrstrcet and Heard, 1982!,

instantaneous annual total mortality Z! of reddrum in Texas bays approximate 1.4 to 1.9. An esti-mate of ! .6 by Matlock �984! was based upon catchcurve analysis and agrees with reported values of Z= 1.39 and 1.90+ 0,13 based on tagging Matlock andWeaver, !979a; Green et al., 1985!. However, theseestimates appear extrcincly high for such an appar-ently long-lived fish, based on theoretical estirnatcsof M �.30.5! using the relationshipbetween mortal-ity and maximu m age Royce, 1972!. Catch curve andtagging data analysis may overestimate Z if there is:I ! hand ling mortality of tagged fish or tag loss; 2! gea rs< lection in which the mesh becomes less and lesscapable ot capturing fish as they grow in size; or 3!change in availability due to seasonal emigrationaway from shorelines in bays or an emigration to thegulf as the sub-adults mature.

Handling inortality is probably minimal bc<ausetagged fish were handled carefully and Elarn �971!found mean mortality of only 6.8 percent for taggedsub-adults held in ponds for 35-424 days. Tag losswas <13 Percent. Together, these sources of error mayadd up to 7 to 20 percent underestimate of Z.

Gear selection likely occurred because trammelnets are selective. Its magnitude is unknown, butrecruitmcnt of sub-adults was knife-cdgcd c.g. oc-curred in a few months!, the effects of which wouldanchor th« left end of a catch curve and emphasizegear selection of larger fish Matlock, '! 984!. Changein a va ilability due to emigration must ha ve occurred.its magnitude a!so is not known, but ]00 percent ofthe sub-adults emigrate. to the Gulf when they reachsexual maturity; they must spend two to four years assub-adults assuming they reach sexual maturity in hrcc to five years, and spend one year as juveniles.

Mrrtfock � Life History 17

Their average anmial change in avai!ability Z! dueto emigration to thc Gulf can be estimated usingRoyce's �972! approach, substituting the duration nfthe sub-adult stage for life span; doing so, avcragcannual values of 2 are �.92-1.53. This range of valuesapproximates values of Z estiinated via catch curveanalysis and suggests simple einigration to the gulfcould account for much of the large va lues of Z foundby catch curve analysis if emigration occurred con-stantly for three to five years. This would be true fnrany catch curve analysis, whether the fish were taggednr not, assuming they emigrate equally and continu-ously to the Gulf. However, emigration is knife-edged, not continuous. Therefore, Royce's �972! ap-proach probably yields gross underestimates of Z inbays when maximum agc instead of maximum timespe~t in bays is uscxl.

Although emigration to the Culf occurs, fishingmay cause most mortality of sub-adults in bays.Based on tag recaptures, at least l7 perccr< r of the sub-adults are caught by fishermen in bays within oneyear after release Green et al., 1985!. However, this isa gross underestimate of bay tag rccapturcs becauseonly 29 percent of the recaptures get reported byrccrea tional fishcrmcn Ma tlock, 1981! . Nonrc port in gmay even be higher than 29 percent for cornrncrcialfishermen because they intentionally discard tags Ma lock, 1984!.

Red drum are quite susceptible to fishing. Theirschooling and tailing behavior may make them easilylocated and captured. Reported tag recaptures inFlorida have been over 47 pcrccnt when rewards of$10,000 were given for returned tags Beaumariage,1969!.

Additional Research

All five gulf states, several universities, and thefederal government, including National Marine Fish-crics Service arc conductmg additional rc~carch onthe life history of red drum in the Gulf of Mexico.Several private companies are also involved in aqua-culture research of red drum, Much information hasbeen generated since 1984 to improve the life historyparameter estimates, but thc basic life history de-scription presented herein remains unchanged. Thereader is refcrr cd to the U.S. Secretary of Commerce'sRed Drum Fishery Management Plan for additionalin forma t ion.

Personal CommunicationsMr. John Brucks, National Marine Fisheries Service,

National Space Technology Laboratory, NSTL Station, Miss.,pers. commun., February 1983.

Mr. C.E�Bryan, Texas Parks and Wildlife Department,Austin, Tex., pers commun., April 1983,

Dr, Mark E. Chittcnden, Jr., Texas AkM University,Collcgc Station, Tex�peri, commun., April 1983.

Ms. Joanne L, Larochc, Gulf Coait Research Labora-tory, Ocean Springs, Miss., pers. commun., April 1983.

Literature Cited

Adkini, C., J. Tarncr, I'. Bowman, and B, Savoie. 1979.A study <>I conimcrcial finfish in coastal Louisiana. La.Dep. Wild!. Fish,, Tech. Bull. 29;62.70.

Amos, A.F. 19HI Redfish kill in thc Gulioi Mexico:June-July 19HI, R/V KATY CRUISE REDFISH- � 1, hydro-graphi< observations. Univ. Tcx. Mar. S<.i. Iriit., Port Aran-iai, I'exai. I Unpublished manuscript!.

Anonym<>ui. 1982. Texas Parki and Wildlile Dcpart-merit, oaita! Fisheries Plan, FY 1982 progr<. si and FY 1983pl,iri PWT i Report 3000-74, 34 pp.

1983. Saltwater finfish research and man-agcmcni in Texas, a report to the Governor and the 68thLegislaturir. Tcx. Parks Wild!. Dep., I'WD Rcport 3000-154,48 pp,

Armstrong, R., M. Devine, J.A. IVIartin, R.F Temple, A,Vastro, C Warsh and R. Whitaker. 1976. >u W.B. Jackson editor!, Fnvironmcntal Studies of the South Texas Con-tinental Shelf, Vol. II. Physical oceanography. NOAAFinal ReFx>rr io Bur, Land Management, In tera gcncy Agree-meni No ilH550-IAS-I9, 290 pp

A.mold, C.A., C.W. Caillouet, Jr., M E. Cliittcnden, Jr.,!.H. Finu<.-anc, R. Juhl, J.A. Martin, !.D. M<'Eachran, E.L.Nakamura, E.T. Park, R.F. Templeand W.L. Trent.1976. p.1425. frr: W.B. Jackson editor!, Environmental Studies ofthe South Texas Outer Continental Shelf 1975, Vol. 1.Plankton and fisheries. NOAA Fina! Rep. to Bur. LandMngmnt., Interagency Agreement No. 0855 l-lA5-19, 425pp

Arnold, C.R., J.D. Williams, A. Johr>s<>n, W.H. Baileyand J.L Lasswell. 1977. Laboratory spawning and larvalraring of red drum and southern flounder, Proc. Annu.Conf. S.E. Assoc. Fish Wild 1. Agencies. 3!:437441,

Bass, R J. and J,W. Avault, Jr. '1975. Food h.>bits, length-wcight relationships, condition factor,and growth of juve-ni!e rcd drum, Scrae>t<>ps ocellata, in Louisiana. Trans, Am.Fish. Soc. 104:3545,

B<wumariagc, D.S. 1964. Returns from thc I 963 Sch!itztagging program. Fla. Board Conser. Tech. Ser. 43, 38 pp.

l 969, Returns from the1965Sch!i tz taggingprogr am i riel uding a cumulative analysis of perv x>us results.F!a. Dep. Nat. Resour. Mar. Lab. Techy. Ser. 59, 38 p.

Beaum.>riage, D.S, and A.C. Wittich. 1966. Returnsfroni the1964 Schlitz tagging program. FIa. Board Conser,Tech. Ser. N<>. 47,51 p.

Bcrc, R. 1936. Parasitic copep<xls from Gu!f of Mexicofish, Ain, MidI. Nat. 17�!;577-625.

I!igelow, H.B. and W.C. Schroeder, l 954. Fiihcs of theGull <>f Maine, U.S. Fish Wildl. Serv. Fish. Bul!. 53�4!:1-s77.

l3oothby, R.N, and J.W. Avault, Jr. 1971. Food habits,length-weight relationships, and condition factorof the reddrum, Sciae>><>ps <>cellata, in southern Louisiana, Trans. Am.Fish. Soc. 100:290-295.

Bowden, K.F. 1967. Circulation and diftusion, p.15-36.in: G.H. Lauff editor!, Estuaries, Amer. Asso. Advanc. Sci.Publ. H3, Washington, D.C,

Bradley, E. and H, Compton.!963. Survey of larval andpoit-larval finfish in Aransasand Corpui Christi Channelsand in thc inshore Gulf of Mexico. Tex. Game Fish Comm.,I'roj. Rep. I961-1962, I'roj. No. MF- R4, 7 p.

Breuer, J.I'., R.L. Benefield, M.G, Wcix<!man, A.R.Martinez, and I, Nava. 1977. Survey of finfish harvest in~elected Texas bays, segment Il. Tex. I'arks. Wildl. Dcp.,Coast. Fish. Rep. Proj. No. 2-231-R-2, 116 pp.

Red Dru>n Aquaeu lture

Briggs,J.C.1974,Marinezoogeography. McGraw-Hill,N.Y., 475 p.

Brown, L.F�Jr., R,A, Horton, J.H. McGowen, C.W.Krcitler, and W.L. Fisher, 1974, Natural hazards of thcTexas coastal zone. Bur. Econ. Geol. Univ. Tex., Austin,1'ex, 13 pp.

Bryan, C,E, 1971. An ecological survey of thc ArroyoCok>redo, Texas, 1966, 169. Tex, Parks Wildl. Dep., Tech.Scr. 10, 28 pp.

Bryan, C.E., T.J. Cody, and G.C. Matlock. 1982. Organ-isms captured by the commercial shrimp f!cet on the Texasbrown shrimp Penaeus aztecus Ives! grounds. Tcx. ParksWil dl. Dep�Tech, Ser. No 31, 26 pp.

Caldwell, K,D and M,C. Caldwell. 1972. The world ofthe bottlenosed dolphin. J.l3, Lippcncott Co., Phila. andN.Y., 157 pp.

Cardcilhac, P.T., C.F. Simpson, F.H. White, N,l'. Th-ornpsi>n, and W.E, Carr. 1981. Evidence ior metal poison-ing in acutcdcathsof large red drum Sciaenops ocellata. Bull,Environm. Contam. Toxicol. 27;639-644.

Carr, W.E.S. and T.B. Chancy. 1976. Harness for attach-ment of an u ltrasonic transmitter to the red drum, Sciaenopsoceilata. Fish. Bull, U.S. 74�!:998,

Causey, D. 1953. Parasitic copepoda of Texas coastalFishc ries, Pub, Inst, Mar. Sci., Univ. Tex. 3�!:5.16.

Chandler, A.C. 1935. Parasites of fishes in GalvestonBay, Proc, U,S. Nat. Mus. 83�977!:123.157.

1954, Ncmatoda. p. 357.358. In: P. Gal tsoff coordinator!, Gulf of Mexico, its origin, waters, and ma-rine life. U.S. Fish. Wild I. Serv., Fish. Bull. 55.

Choo, L,N, 1978. A basis tor classifying western Atlan-tic Sciaenidae Teleostei: Perciformes!. U.S. Dep. Commer�NOAA Tech, Rep. NIVIFS Circ. 415, 64 pp,

Chao, L.N. and J.A. Musick. 1977. Life history, fccxiinghabits, and I'unctional morphology of juvenile sciaenidfishe~ in thc York River Estuary, Virginia. Fish. Bull, U.S.75�!:657-702.

Chittcnden, M.E.Jr. and D. Hoore. 1977. Compositionof the ichthyotauna inhabiting the 110-mctcr bathymetriccontour of the Gulf of Mexico, Mississippi River to the RioGrande. N.E. Gu lf Sci, 1�!:1 i�-114.

Chittcnden, M,E�Jr. and J.D. McEachran. 1976. Com-position, ecology, and dynamics of demersal fish com-munities on the Northwestern Gulf of Mexico continen-tal shelf, with a similar synopsis for the entire Gulf. ScaGrant Publ. TAMU-SG-76-208, 104 pp.

Christmas, J.Y., G. Guntcr, and E.L. Whatley. 1960.Fishes taken in the menhaden fishery of Alabama, Missis-sippi, and eastern Louisiana, U.S. Dep. Interior,Fish WildlSer., Spec, Sci, Rep;Fish. No. 339, 10 pp.

Cody, T.J. and R.H. Avent. 1980, Shrimp and Finfishinvestigations in the North wcstcrn Gulf of Mexico. Part III.Assessment of bottom longline fishing off the central Texascoast. Tex. Parks. Wildl. Dcp., Coast. Fish. Branch, Mng-mnt. Data Ser., 24 p. Unpublished manuscript!.

Cody, T.J., B.E. Fuls, C.C. Matlock, and C.E, Bryan,1981. Assessment of bottom longline fishing off the centralTexas coast, a completion report. Tex. Parks Wild l. Dep.,Coast. Fish. Branch, Mngmnt. Data Ser, 22,51 pp.

Cody, T.J., K.W. Rice, and C.E. Bryan. 1977. Foodfishand potential foodfish captured in a large-mesh fish trawlalong the central Texas coast. Tcx. Parks Wild 1. Dep�Coast.Fish, Branch, 49 p. Unpublished manuscript!.

Colura, R. and B. Hysmith.1976. Fingcrlmg productionof spotted seatrout, Cynoscion nebulosus, and red drum,Sciaenops ocellata, in saltwater ponds. Annu. Rep. Mar,

Fish, Res, Sta,, Tex. Parks Wildl. Dep�Palacios, Tcx., 39pp

Colure, R I, 8 T Hysmith and R.E, Stevens.1976. Fin-gerling product ion of stri pcM bass Moro ne saxatr'lis!,spot tedseatrout < ynoscirn nebulosus! and red drum Sciaenopsocellata! in saltw.>ttr ponds. Proc. 7th World Maricult.Soc,p. 79-92.

C<>mpton, H. 1965. Survey of the fishes found in theinshore Gulf of Mexico and ol post-larval fishes inAransas,Pi>rt Mansfield and Port Isabel Ship Channels. Tex, FishGame Comm., Proj. Rep., Proj. No. MR-R-6, p 383-413.

Compti>n, I L and E. Bradley. 1964. A study of the post-larval pcnacid shrimp entering Aransas Bay. Tex, ParksWildl. Dep., Coast, Fish, Branch, Proj. Rep. 1962-1963, p.127-143,

Crocker, P.A., C R. Arnold, J.A. DeBoer, and J.D.. Holt.1981. Preliminary evaluation of survival and growth ofjuvenile red drum Sciaenops ocellata! in fresh water. J.World Maricult, Soc, 12<1!:122.134,

Crocker, R,A. 1965. Planktonic fish eggs and larvae ofSandy Hook estuary. Chesapeake Sci. 6�!:92-96,

Darnell, R.M. 1958. Food habits of fishes and largerinvertebrates ot Lake Pontchartrain, Louisiana, an estuar-inccommunity. Publ. Inst. Mar. Sci., Univ, Texas. 5:353416.

Diener, R.A., A. Inglis and G, B. Adams. 1974. Stomachcontents of fishes from Clear Lake and tributary waters, aTexas estuarine area. Contrib. Mar. Sci. 18;i-li,

Dokken, Q.R. 1981. Spatial and temporal distributionand species comp>sition of larval fish populations withinAlazan Bay, Texas. M.S. Thesis, Corpus Christi State Univ.,61 pp.

Dove!, W. 1967. Fish eggs and larvae of the MagoihyRiver, Maryland. Chesapeake Sci. 8�! l25-129.

Dugas, R,, V. Cuillory and M. Fisher. 1979. Oil rigs andoffshore sport fishing in Louisiana. Fisheries. 4<6!:2-10.

Donham, F. 1972. A study of commercially importantcstuarinc-dependent industrial fishes. La. Wildl. Fish.Comm., Tech. Bull. 4, 63 pp.

Elam, L.'!971. Fvaluation of fish tagging methods Tex,Parks Wild 1. Dep., Coast. Fish. Proj. Rep., p. 39-55.

Finucanc, J.H., L.A. Collins and J.D. McEachran. 1977,lri: W.B. Jackson editor!, Environmental studies of theSouth Texas outcr continental shelf, ichthyoplankton-mack-erel eggs and larvae. NOAA Final Rep. Bur. Land Mngmnt,Interagency Agreement No. AA550-1A7-3, 484 pp.

Fontenot, B.J., Jr. and H.E. Rogilio, 1970. A study ofestuarine sport fishes in the Biloxi marsh complex L.ouisi-ana. L. Wild!. Fish. Comm., D.J. Proj. F-8 Comp!. Rep., 172pp.

Green, A.W., H.R, Osburn, G.C. Matlock and H. E.Hegen. 1985. Estimated survival rates for red drum inNorthwest Gulf of Mexico bays. Fish. Res. 3;263-277.

Guest, W,C. and J.L. Lasswcll. 1978. A note on court-ship behavior and sound production of red drum. Copeia.1978�!:337-338.

Guntcr, G. 194 I. Death of fishes due to cold on thc Texascoast, January, 1940, Ecology. 22:203-208.

Gu nter, G. 1942. Contributions to the natural history ofthc bot tlenosc dolphins, Tursiops truncatus Montaguc!, onthe Texas coast, with particular reference to food habits. J.Mamm. 23�!:267-276

Guntcr, G. 1945. Studies on marine fishes of Texas.Publ. Inst. Mar. Sci., Univ. Tex. 1�!:1-190.

Gu nter, G. 1952. Thc import of catastrophic mortalitiesfor marine fisheries along the Texas coast.J. Weld l. Mngmnt.16�!:1-7.

Afatlock � Life History19

Gunter, G. 1958. Population studies of the shallowwater fishes of an outer beach in South Texas. Pub!. Inst.Mar. Sci., Univ. Tex. 5:186-193.

Guntcr, G, and H.H, Hildcbrand. 1951. Destruction offishes and other organisms on the south Texas coast by theco!d wave of January 28-February 3, 1951. Ecology 32:731-735

Hamilton, C.L. 1980. Texas commercial harvest statis-tics, 1978-79. Tex. Parks Wi!dl. Dcp., Coast. Fish. Branch,Mngmnt. Data Scr. 12, 32 pp.

1981. Texas commercial harvest statistics,!979-80. Texas I'arks Wild!. Dep., Coast. Fish. Branch,Mngmnt. Data Scr, 26, 42 pp,

1982. Texas commercial harvest statistics,1977-81. Tex. Parks Wild!. Dep., Coast. Fish. Branch,Mngmnt. Data Scr.45, 59 pp.

Harden Jones, F.R, 1968. Fish migration. Edward Amo! dLtd., London, 325 pp.

Harrington, R. 1973. Fish graveyard no longer trapsvictims. Tex. Parks Wildl. Magazine, July:16-17.

Heftcrnan, T.L., A.W. Grccn, L.W. McEachron, M.G.Wcixclman, P.C. Hammcrschmidt and R.A. Harrington.1977. Survey ot finfish harvest m se!ected Texas bays, Tex.Parks Wi! d I, Dcp,, Coast, Fish, 8 ra nch Proj, No, 2-231-R-I,I16 pp.

Heffernan, T.L. and R.J. Kemp. 1980. Management ofred drum resources. Proc. CoHoquiumonthe Biology andManagementof Red Drum and Seatrout, Gulf States Mar.Fish. Comm., Oct. 19-21, 1978, Tampa, Fla,

Hegen, H,E. 1981. Monitoring of coastal finfish re-sources for sport fish management, October 1979-Septem-ber 1980, Tex, Parks Wildl. Dep., Coast. Fish. Branch,Mngmnt. Data Ser. 28, 228 pp.

1982. Monitoringofcoastal finfish resourcesfor sport fish management, October 1980 September 'I 981.Tex. Parks Wildl. Dep., Coast. Fish. Branch, Mngmnt.Data Ser. 33, 186 pp.

Hegen, H.E. and G.C. Matlock. 1980. Monitoring ofcoastal finfish resources for sport fish managmcnt, October1978-September 1979, Tex. Parks Wildl. Dep., Coast. Fish.Branch, Mngmnt. Data Ser. ! 7, 245 pp.

Hcin, S.H. and J.A. Shepard. 1980, Growth of juvenilered d rum, S ciaenops mllata, in a quarter-acre pond. Irt: Con-tributions of the Marine Re~rch Laboratory-1978, Ia,Dcp. Wildl. Fish. Mar. Res, Lab., Grand Terre Island, La.,Tech. Bu ll. 31, 86 pp.

Hen! cy, H,W. and D.H. Lewis. 1976. Anaerobic bacteriaassociated with epozootics in grey mullet <1Vfugtl cephatus!and red fish Sciaenops ocellata! along the Texas Gulf Coast.J. Wild l. D is. 12:448453.

Hermann, S.S. 1963. Planktonic fish eggs and larvae ofNarragansett Bay. Limnology Oceanog, 8�!:103-109.

Hildebrand, H.H, 1954. A study of the brown shrimp Pertaeus aztecus Ives! grounds in the Western Gu! f of Mcxim.Pub!. Inst. Mar. Sci., Univ. Tex. 3�!:1-366,

Hi!debrand, S.F. and L.E, Cable, 1934, Reproductionand development ot whitings or kingfishes, drum, spot,croaker and weakfishes or sca trouts of the United States.Bu! I. U.S. Bur. Fish. 48<16!:41-117.

Hildebrand, S.F. and W.C. Schroeder. 1928. Fishes ofChcsapcake Bay. Bull, U,S. Bur. Fish. XLIII:1-388.

Hoese, H.D. 1965. Spawning of marine fishes in the PortAransas, Texas area as determined by the distribution ofyoung larvae. Ph.D. Dissertation, Univ. Tex., 144 pp.

Holland, J S, N,J. Macio!ck, R,D, Ka!ke and C.H.Oppenheimer. 1973. A benthos and planktonstudy of thcCorpus Christi, Copano and Aransas Bay systems. I, Re-port on the methods used and data collected during thcperiod September 1972-June 1973. Unpub!. 1st Annu. Rep,to Texas Water Dove!op. Board, Univ. Tex, Mar, Sci. Inst.I'ort Aransas, Tex. 118 pp.

1974. A benthos and plankton study of theCorpus Christi, Copano, and Aransas Bay systems, II.Report on data collected during the period July 1973-April 1974. Unpubl. 2nd Annu. Rep. to Tcx. Water De-velop. Board, Univ. Tex. Mar, Sci. Inst., Port Aransas, Tcx.,121 pp.

Holt, ]., G. Godbout and C.R. Arnold. '! 981a, Effect oftemperature and salinity on egg hatching and larval sur-viva! of red drum, Sciaenops ocellata. Fish. Bull., U.S.79�! 569-573.

Holt, J., A.G. Johnson, C,R, Arnold, W.A. Fable,Jr. andT.D. Wi!!iams 198]b. Description of eggs and larvae oflaboratory reared red drum, Sciaenops ocellata. Copeia.I 981:751-756.

Holt, S.A�C,L. Kitting and C.R. Arnold. 1983. Distribu-tion of young red drums among different sea-grassmeadows. Trans. Am. Fish. Soc. 112:267-271.

Houde, E,D., J.C. Leak, C.E. Dowd, S.A. Bcrkley andW.J. Richards, 1979, Ichthyoplankton abundance and di-versity in the eastern Gulf of Mexico. Rcp. Bur. LandMngmnt., Contract No. AA550-CT7-28, U.S. Dep. Com-mcr. Nat. Tech. Info. Serv., PB-299839, 546 pp.

Hysmith, B.T., R.L. Colura and G.C. Mat!ock, 1983,Effects of stocking rate and food type on growth andsurvival of lmger!ing rcd drum. Proc. Warmwater Fish.Cult, Workshop, Annu. Meet. World Maricult. Soc., March14, 1982, Charleston, SC in press!.

Ingle, R.M,, R.F. Hutton and R.W. Topp. 1962, Resultsof the tagging of saltwater fishes in Florida. Fla, BoardConsev. Tech. Ser. 38,57pp.

Ivcrscn, E.S. and B. Yokel. 1963. A myxosporidian<sporozoan! parasite in the red drum. Bull, Mar. Sci. GulfCaribb. 18:449-453.

Jannke, T.E. 1971. Abundance oi young sciacnid fishesin Evcrg!adcs National Park, F!orida, in relation to seasona nd other variables. Univ. of Miami Sea G rant Tech, Bull.11, 128 pp.

Johnson, A.G., W.A. Fable, Jr., T.D. Williams and C. R.Arnold. 1977. Description of reared eggs and young larvaeof the red drum, Sciaenops ocellata. p. 118-140. lrn MarineFish Propagation Study, Fed. Aid Proj. I-31-R, Comp!.Rep., Tex. Parks Wi!dl. Dep.

Juneau, C.L., Jr. 1975. An inventory and study of thcVermi! ion Bay-Atchafalaya Bay complex, Phase II; biology.p21-67. In. An inventory and study of the Vermilion Bay-Atchafalaya Bay complex. La. Wildl. Fish. Comm., Tech.Bull. 13.

Kemp, R.J. 1949. Report on stomach analysis on June 1,!949 through August 31, 1979, Tcx. Game, Fish, OysterComm., Mar. Lab. Annu. Rep. �948-1949!:101-127.

Kiiby, J.D. 1955. The Fishes of two Gulf' coastal marshareas ot Florida. Tulane Study. Zool. 2<8!:176-247,

King, B D., ll!,1971.Study ofrnigratory patterns of Ashand shellfish through a natural pass. Tex. Parks Wildl,Dep. Tech, Ser.9,54 pp,

Kna pp, F, T. 1950. Menhaden utilization in relation tothe conservation of food and game fishes of the Texas Gulf

20 Red Drum Aquaculture

coast Trans, Am. Fish. Soc. 79:137-144,Linton, E, l901. Parasites of fishes of thc Woods Hole

Region. Fish. Hull., U.S. 19:405-492.1905. Parasites of fishes of the Beaufort,

North Carolina region. Fish. Bull, U.S. 24:323-428Lippson, A.J, and R.L. Moran, 1974. Manual for iden-

tification of early developmental stages of fishes of thePotomac River Estuary, Env. Tech. Center, Martin Mari-etta Corp., Baltimore, MD., 282 pp.

Loman, M.1978.0therfinfish. p.143-147. In J.Y Christ-mas editor!, Fisheries assessment and monitoring, Mis-sissippi. P.L. 88-309,2-215-R,Compl. Rcp., Gulf Coast ResLab., Ocean Spiingvs Miss.

Luebke, R,W. and K. Strawn, 1973. The growth, sur-vival, and feeding behavior of rcdfish Sciaenops ocellata! inponds receiving heated discharge water from a powerplant, Proc,4th Annu. Workshop World Maricult. Soc., p,'143-154.

McEachron, L.W. and G.C. Matirick, 1983. Harvestestimate of the Texas Chartered boat fishery. 1VIar. Fish.Rev. 45�!;1 I-1'7.

Mc garland, W. 1959. Distribution and infestation of theparasite Poerilancistrium robustrum in some Texas coastfishes. Tcx, Game Fish Comm. Proj. Rep. ]958-1959!:11pp.

McFa rland, W.N. 1963, Seasonal change in the numberand biomass of fishes from the surf at Mustang island, Tex.I'ubl. Inst. Mar, Sci., Univ. Tex. 9:91.105.

McKee, D.A, 1980. A comparison of the growth rate,standard length-weight relationships and condition factorof red drum, Sciaenops ocellata Linnaeus!, from an electricgenerating station's cooling lake and the natural cnviron-mcnt. M.S.Thesis, Corpus Christi State Univ�53 pp.

Mansu eti, R.J. 1960. Restriction of very young rcd drum,Sciaenops ocellata, to shallow estuarine waters of Chesap-eake Bay during late autumn. Chesapeake Sci. 1:207-210,

Marley,R.D 1983.Spatialdistributionpatternsofplank-tonic fish eggs in lower Mobile Hay, Alabama. Trans. Am.Fish. Soc. 112:257-266.

Massrnann, W,H,, J.J. Norcross and E.H, Joseph. 1961,Fishes and fish larvae collected from Atlantic planktoncruises of R/V PATHFINDER, December 1959-December1 960. Va, Inst. FIar. Sci,, Va. Fish, Lab., Spec. Sci, Rep. 26,22 pp,

Massrnann, W. H., E.B. Joseph and J.J. Norcross. 1962.Fishes and fish larvae collected from Atlantic planktoncruises of R/V PATHFINDER, March 1961-March 1962.Va, Inst. Mar. Sci., Spec, Sci. Rep. 33, 23 pp.

Matlock, G.C, 1979. The use of fish trawl to sample largefish. Tex.'Wild L Dep., Coast. Fish. Branch, Mngmnt. DataSer. 2, pp.

1980. History and management of the reddrum fishery, Proc. Colloquium on the Biology andManagement of Red Drum and Seatrout, Gulf States Mar.Fish. Comm., Atlantic States Mar. Fish. Comm.,Oct. 19-21,1978, Tampa, Fla.

Matlock, G.C. 1981. Nonreporting of re apturcd taggedfish by saltwater recreational boat anglers in Texas, Trans.Am. Fish. Soc. 110:90.92.

1982, Thc conflict between user groups ofred drum and spotted seatrout in Texas. p. �1-108. ln: R.H.Stroud cditor!, Marine Recreational Fisheries, Proc. 7thAnnu. Mar. Rec. Fish. Symposium, Fort Lauderdale, Fla.,May 10-11, 1982.

1983, Angler origin and retention rateinterpretation. N. Am, J. Fish. Mngmnt. 3;438-444.

Matlock, G.C. 1984. A basis for the development of a

management plan for red drum in Texas, I'h,D, Disserta-tion, Texas ARM L niv., College Station, Tex. 29I pp.

Mat lock, G.C. 1986. Rcd drum sex ratio and sire atsexual maturity, Tex. Parks Wildi. Dep,, Coast. Fish. BranchMngmnt. Data Ser. No. 85,6 pp.

Matlock, G.C. and l.E. Weaver.1979a. Assessment andmonitoring of Texas coasfai finfish resources. Tcx. I'arksWildl. Dep., Coast. Fish. Branch, Rep. Proj. No,2-313-R,247pp

19 9b. Fish tagging in Texas bays LluringNovember 1976-September 1976. Tex, Parks Wildl. Dep.,Coast. Fish. Branch, Mngmnt. Data Ser. No. 1, 136 pp.

Mat lock, G,C., RJ. Kemp,Jr. and T.L. Hcffcrnan.1986.Stocking as a management tool for a red drum fishery, apreliminary evaluation. Tex. Parks Wildl. Dept., Coast.Fish. Branch, Mngrnnt. Data Ser. No. 75, 27 pp.

Matlock, G.C.,J E. Weaver, L.W. McEachron,J.A. Dailcy,P,C. Hafnmcrschmidt, H.E. Hegcn, R.A. Harrington andG.M. Stokes 1978. Trends in finfish abundance in lexasestuaries as indicated by gill nets, Tcx, Parks Wildl. Dep.,Coast. Fish., 271 pp.

Miles, D.W. 1950. The life histories of the spottedscatrouf. Cynoscion nebulosus, and the redfish, Snaneopsocellatus. Annu. Rcp. Tex, Came, Fish,Oyster Comm., Mar.Lab., p. 1-38.

lvlilc~, D.W. 1951. Thc life histories of the scatrout,Cynoscion nebulosus, and thc redfish, Sciaenops ocellatus,sexual dcve!opment Tcx. Game, Fish, Oyster Comm., Annu.Rcp. 1 September 1950-31 August 1951, 1 pp.

Miller, G.L. and S.C, Jorgcnson. 1973. Mcristic charac-ters of some marine fishes of the wcstcrn Atlantic ocean.Fish. Bull�U.S. 71:301-312.

Miller, J.M. 1965. A trawl survey of thc shallow Gulffishes near Port Aransas, Texas. Publ, Inst. Mar, Sci., Univ.Tex. 10:80-107,

Moddc, T. and S.T. Ross. 1981. Seasonality of fishesoccupying a surf xone habitat in thc northern Gulf ofMexico. Fish. Bull., U.S. 78:911-922.

Moe, M A, Jr. 1972. Movement and migration of southRorida fishers. Fla, Dcp. Nat. Rcsour. Tech, Ser. 69, 25 pp.

Moffett, A,W., L.W, McEachran and J.G. Key. 1979.Observations on thc biology of sand seatrout Cynoscionarenarfus! in Galveston and Trinity Bays, Texas. Contrib.Mar. Sci. 22:! 63-172.

Murphy, M. and M,E. Chittcnden, Jr. 1982. Aspects ofthe life history of the Gulf butterfish Prdus burti! <Unpub-lished manuscript!.

Neff, J N., M,P. Cogliansc, L.A. Reitscma, S. Andersonand W, McCullouch. 1981. Brine toxicity bioassays onrcdiish. Vol V Pari A!. Ini W.B. Jackson editor!, Shrimpand redfish studies; Bryan Mound brine disposal site offFreeport, Texas, 1979-1981. NOAA Tech. Mem<irandumNMfS-SEFC-69, 82 pp.

Odum, W.E. 1971. Pathways of energy fhow in a cmouthFlorida estuary. Univ. Miami Sea Grant Tech. Bull. 7, 162pp.

Osburn, H,K�G.C. Matlock and A,W. Green, l982. RedDrum Sciaenops ocellatus! movement in Texas bays. Cont.Mar. Sci., Univ. Tex. 24:85-97,

Overstrcct, R.M.1977. Poecilancfstriumcaryophyllunf andother trypa norynch ccstode plcrocercoids from the rnu scu-laturc of Cynoscfon nebulosus and other sciaenid fishes in thcGu li of Mexico. J, Parasitoi. 63�!;780-789.

Overst rect, R.M. and R.W. Heard. 1978. Food of thc rcddrum Sciaenops ocehata from the Mississippi Sound, GulfRes. Rep, 6�!:13'1 -1 35.

1982 Food contents of six commercial fishes

Mattock � Life H>'sfory 2]

from Mississippi Sound Cu! f Res. Rep. 7;137-15kOwens, DW.,K,A.Jones and BJ. Ga!! oway. 1981. Brine

avoidance/attraction bioassays on redfish. Vo!. V part 13!.ln: W.H. Jackson editor!, Shrimp and rcdfish studies;Bryan Mound Brine disposa! sites off Freeport, Texas,19'79-1981, NOAA Tech. Memorandum NMFS-SEFC+9,»pp

Pearse, A.S. 1952. Parasitic crustacea from thc Texascoa~t. Publ. Inst. Mar, Sci., Univ, Tex 2�!;5-43.

Pcarson, J.C. ]929. Natural history and conservation ofthe redfish and other commercial sciaenids on the Texa~coast. Hull. Bur. Fish. 44;]29-214.

!94]. The young of some marine fishestaken in lower Chesapeake Hay, Virginia, with spccia!reference to the gray sca trout, Cynoscion regafis H!och!.U.S. Dep. Interior, Fish Wildl. Ser., Fish Bull. 50:79.102.

Pcrret t, W S,,J,E, Weaver, RO. Williams, PL. Johansen,T.D. Mcl!wain, R.C. Rau!erson and W.M, Tatum. !980.Fishery profiles of red drum and spotted seatrout. GulfStates Mar, Fish. Comm,6,60pp.

Powles, H. and B.W. Stcnder. 1978. Taxonomic data onthc carly life history stages of Sciaenidae of the SouthAtlantic Bight of the United States. South Carolina Mar.Res. Center Tech. Rep. 31 pp,

Rabelais, S.C,, C,R. Arnold and N.S, Wohlschlag, ]981.Thc cFfects of IXTOC I oi] on thc eggs and larvae of reddrum Sciaenops ocellata!. Tex. J. Sci. 33:33-38.

Reid, G.K,,Jr. ]955. A summer study ot the biology andecology of East Hay, Texas. Part Il, Thc fish fauna of Eastbay, the Gulfbeach, and summary. Tex. J. Sci. 7�!:430453.

Reid, G,K., A. Inglis and H.D. Hoese. 1956. Summerfoods of some fish species in East Hay, Texas. SouthwestNat. 1�!:100- 104,

Riggin, G.I., Jr. and A.K. Sparks. 1962. A newgasterostome, B>icephalo>des rnegac>rrus, from the redfishSciaenops ocella ta, Proc. Helm, Soc. 29;27-29.

Roberts, D.E., Jr�B,V. Harpstcr and G.E. Henderson.1978a, Conditioning and induced spawning of the reddrum Sciaenops ocellata!. Proc. Ninth Annu. Meet. WorldMaricuit, Soc., p. 333-343.

Roberts, D.E., Jr,, L.A. Norey, III, G.E. Henderson andK.R. Ha!scott. ]978b. The effects of delayed feeding, stock-ing density and food density on survival, growth andprod uctii>n of larval red drum Sciaenops ocellata. Proc. NinthAnnu. Meet World Maricult. Soc., p.333-343,

Roess!er, M. 1967. Observations on the seasonal occur-rence and life histories of fishes in Huttonwood Canal,Everglades National park, Fla. Ph.D. Dissertation, Univ. ofMiami, ! 55 p.

Rohr, H.A, 1968. SurFacc schooling of red drum. Under-water natur. 5�!:21-23.

Rohr, H.A. 1980. Use of hard parts to age Gu!f of Mexicored drum, Sciaenops ocellata. Proc. Colloquium on the Biol-ogy and Management of Red Drum and Seatrout, p. ,

Ross, J.L,, F.S. Pave!a and M.E. Chittenden, Jr. 1982.Seasonal occurrence of black drum, Pogonias cromis, androd drum, Sciaenops ocellat»s, off Texas. N.E. Gulf Sci. Inpress!.

Rounsfcll, G,A. 1975. Ecology, utilization and man-agement of marine fishes. C.V. Mosby Co., St. Louis, No.,516 pp.

Royce, W.F. 1972. Introduction to the fishery sciences.Academic Press, N.Y.,351 p.

Sawyer, R.T., A.R, Lawlcr and R.M. Overstreet. 1975.Marine!ocxhes of the eastern United States and thc Gulf ofMexico with a kcy to the species. J. Nat, Hist. 9:633-667.

Shlossman, P.A. and M.E. Chittcnden, Jr. 1981. Repro-

duction, m<>vc>ncnts,and populationdynamicsofthcsandseatrout, Cynoscron arenar>us, Fish. Bull., U,S. 79:649-669.

Simmon~, E.G. 1957. An ecological survey iif thc upperLaguna Madrc of Texas, Pub!. Inst. Mar. Sci., Univ. Tex.4�!:156-20 !.

Simmons, E G, and J P. Hreucr.]962. A study of redfish,Sciaenops i>cellata Linnaeus,and b!ackdrum, Pogonrascro>nis!.innaeus I'iibl. Inst Mar. Sci., Univ. Tex, 8:184.21],

1982, Fish tagging on the Texas coast, Tex.Parks Wild!. Dept., Coast, Fish. Proj. Rep. 1976, p. 66-107.

Sparks, A,K. !958. Some digenetic trematodes of fishesof Grand Isle, Louisiana, Proc. La. Acad. Sci 8, 20:71-82,

Spr>nger, S. and H.R. Hullis, Jr. 1956. C<>Bections by thcOregon in the Gulfof Mexico. Llfcofcrustaccans, mollusksand fishes identified from collections made by the explora-tory Fi~hing vessel Oregon in the Gulf of Mexico andadjacent seas 1950 through ]955. Spec. Sci. Rep. U.S. Fish,Wild!. Serv. 196:!-34,

Springer, V.G. and K.D. Woodbum. 1960. An cco!ogi-cal study of the fishes of the Tampa Hay Area. Fla. StateHoard Conser., Prof. Paper Ser. 1:104 pp.

Standard, G. and M,E. Chittenden, Jr. 19�4. Reproduc-tion, movements and population dynamics ot thc bandeddrum, far>mus fasciatus, in the Gulf of Mexico. Fish, Hull.82:] -27.

Sykes,j.E.and J,H.Finucane. 1965,0ccurrencein TampaBay, Florida, of immature species doininant in Gulf ofMexico commercial fisheries. Fish. Bull., U.S. 65�!:369-379.

Thicling, D.L. and H.A. Loyacano, Jr. 1976. Age andgrowth of red drum from a saltwater marsh impoundmentin South Carolina. Trans. Am, Fish. Soc, 105:41-44.

Topp, R. !963. The tagging of fishes in Florida, 1962program, Fla. Board Conser., Mar. Res, Lab., Prof. PaperSer. 5:76 pp.

Topp, R.W. and C.F. Cole, 1968. An ostcx>logica! studyof t he sciaenid Genu s, Sciaenops Gill Tc! coster i, Sciaenid ac!.Bu! I. Mar. Sci. 1 8:902-945.

Trimble, W.C. 1979. Yield trials for red d rum in brack-ish-wa ter ponds, 1976-1979. Proc. A nnu. Con F. S.E, Assoc.Fish. Wild!. Agencies. 33:432-44].

Wakefield, C,A. and R,L. Colura, 1983. Age and growthof red drum in three Texas bay systems. Proc, Annu. 1VIeet.Tex. Chap, Am, Fish. Soc., Vol, 5. In press!.

Watson, R.L, and E,W. Bchrens. 1970. Nearshore sur-face currents, south-eastern Gulf coast. Contrib. Mar. Sci.,Univ. Tex. ]5:]33.�4.

Weixe! man, M.G. 1982. The f'all rcd drum Gu! f of Mexicopier fishery off Galveston Bay, Texas, Tcx. Parks Wild!.Coast. Fish. Branch, Mngmnt. Data Ser. 42, 23 pp.

We!sh, W.W. and C.M. Brcder, Jr. 1924. Contributionsof the life histories of Sciacnidae of the eastern UnitedStates coast. Bull. U.S, Bur. Fish. 39:14] -201.

Wilson, C G. 1905, North American parasitic copepodsbelonging to the Caligidae. Proc. U,S. Nat Mus. 28 479-672,

1915. North American parasitic copepodsbelonging to the Lernaropodidac, with a revision of theentire fami!y. Proc. U.S. Nat. Mus. 47;565-729.

1932. The copepods of the Woods Holeregion. Hull. U,S. Nat, Mus. ]58�0!:] 435.

Wohlschlag, D,E. 1981, Natural factors affecting rc-cruitinent. p. 177-'l89. ln: H. Clcppcr editor!, MarineRecreational Fisheries 6. Proc. 6th Annu. Mar, Rec. Fish.Symposium, Houston, Tcx., March 19-20, ]981, 216 pp.

Yokel, H.J. 1966, Contributions on the biology anddistribution of red drum, Sciaenops ocellata. Master's Thesis,Univ. of Miami, 160 pp.

23Sroingte � Status of Fi»4ery

rted commercial and estimated recreational red drum landings thousands of pounof Mexico, 1979-1986.

Rccrca t iona I' Conrmercia I'

Sta te

Wa tcr» EEZ Total'

State

Wa ters EEZ

te

rs' EEZ Total'

80 2,771

48 2,729

31 2,748

77 2,425

206 3,087

987 4,334

3,457 6,343

8,184 13,535

11,227

9,544

8,068

12,607

8,278

8,281

9,098

] 0,223

34 8,570 2,691

1,282 8,145 2,681306 5,657 2,717

475 1�,734 2,348

2,065 7,462 2,881

1,491 6,425 3,347

324 6,536 2,886

232 4,080 5,346

114

1,330 I

337

552

2,271 I

2,478 I

3,781 1

7,268 I

�

52 5,977 53,524 19,551 4,886 24,437 67,103 10,863 7

93 854 7,647 2,793 698 3,49'I 9,580 1,552Kecreational Fishery Statistics Survey data provide to 8 MFS Southeast Fisheries Center by 13.'Source: NMFS Landing Statistics, 1979-19tf5; 1985 data are preliminary.'Landings in state watc

gs for which the area of capture is unknown. " May not eifuaI column totals due to rounding o'Preliminary data subject fo change, Texas data and hcadboaf data not aw ail*bio.' for 1979-19ff5.

tion rates arc and have been higher than the levelwhich v:ould maintain the spawning stock, even ifno offshore fishing occurs on the spawning stock.They have indicated that in order to assure thatoverfishing docs not occur the stock must not be re-cluced below 20 to 40 percent of the spawning stocksize that existed before any fishing occurred.

Therefore, the Gulf-wide status or condition ofthc fishery is not good and rather significant regula-tory actions are and will be required to assure thespawning stock is restored and maintained and toimprove thc productivity of the inshore fishcricw.Thcsc regulatory actions will result in restrictedharvest levels for fishermen in the near-term in orderto increase long-tcrrn productivity from the fishery.

The Gulf states are taking and planning actions toregulate the inshore fishery to allow a minimumescapcmcnt level of 20 percent of the fish that wouldhave escaped had there been no fishery. This meansthat 20 percent of each yearwlass ot fish should beallowed to survive in the inshore fishery so they maymigrate to the offshore spawning stock when theyreach thrcc or four years of agc or approximately 30inches in length. Considering that annual cscape-mcnt levels have 4wn as Iow as one percent of eachyear-class as carly as 1964 for Florida and muchbelow 20 percent for other states more recently, evena tta ining a 20 percent escapement level will rcquircseveral years for some state regulatory agencies.

NMFS thr'ough implementation of a SecretarialFishery Management Plan FMP! and the Council

through an amendment to that plan have taken ac-tion» to prohibit directed harvest of rcd drum fromthe offshore spawning stock, allowing only smallallocations for incidental bycatch front the EEZ byrecreational and cornrncrcial users. Thc Council'samendment will prohibit such directed harvest untilthc escaperncnt goal of 20 percent is reached by thestates. It also prohibits any retention of red drumfrom fcdcral waters off Texas and Florida v here datasuggested the historical levels of escapement to thespawning stock were lowest. Therefore, for theimmediate future, total landingsby recreational andcommercial users from federal waters will not exceed625,000 pounds, annually.

State'

Actions by the states to increase escapement ofjuveniIc fish to the spawning stock will similarlyreduce harvest for inshore waters. 'I' t, xas Parks andWildlife Department, who recognized ovcrfi»hingwa» occurring in their fishery in the mid-'1970s,implemented progressively more restrictive rules toreduce fishing harvest levels. Based on a legisIativcmandate, thev implemented a prohibition on sale ofrcd drum in 1981. Currently their rulc» provide for arecreational bag limit of five fish that must be be-tween 18 and 30 inches in length. Restrictions such as

'State rules have been changing rapidly <n ejforts to increaseescapement of juveniles to 30 percent the current' escapementgoat!. Each state should be contacted for current rates.

Red Drum Aquaculture

these not only will increase cscapcmcnt, but will alsoover tirnc increase the poundage available from theinshore fishery. Texas also has been stocking hatch-ery-reared rcd drum to increase rccruitrnent to thefishery,

Florida has currently closed its red drum fisheryto all harvest by emergency rule tha t will bc extend cdby permanent rule while they develop conservationmeasures to increase the escapemcnt level. Alabamaprohibits sale of rcd drum from its waters and isconsidering a reduced bag limit and size limits fortheir fishery. Mississippi limits commercial catch bya 200,000 pound annual quota and recreational catchby size limits and a bag limit of ten fish. Louisiana'slegislature is considering bills which would imple-mentt commercial quo tas and more restrictive recrea-tional bag and size limits.

The status of the fishery is, therefore, in somewhatof a flux and probably will remain so for severalyears. Fortunately, howcvcr, a great deal of addi-tional scientific information is being collected on thefishery that will give periodic bench marks to betterassess the status of the stock and whether regulatoryrestrictions should be increased or relaxed.

Red Drunr Aquacu ture

Catfish Farming IndustryDevelopment

HatcheryAttempts to determine thc spawning

rcctuirements of channel catfish startedin the carly 1900s Lcary, 1910!. Theseattempts to successfully spa wn channelcatfish werc continued by a number offishery scientists and federal and statefish hatchery workers Clapp, 1929;Mobley, 1931; Lenz, 1947!. By 1950 thebasic methods for spawning channelcatfish had Exxon developed and arc stillin use today. Thus, it took about fivedccadcs to get through the first phaseand only about three and a half decadesto reach a viable, economically impor-tan t industry worth an es tirna ted $300 to$350 million to farmers in I 986,

In addition to learning how to spa wnand hatch eggs, research must be di-rected at the development of the technol-ogy needed to domestic brood stock. Aviable commercial industry can seldomdevelop if brood stock must bc capturedannually from the wild.

Feed

Early a t tempts to raise fingerling andf<xxl-size catfish centered around theuseof inorganic fertilizers to increase thenatural productivity of the ponds andthe usc of a variety of food items tosupplement thc natural food available Morris, 1939; Swingle, 1954!. Maximumannual yields using fertiliza tion only was92 pounds per acre. Feeding soybeancake increased this yield to only 247pounds per acre per year Swingle, 1954!.These production levels were too low toform the basis for a viable commercial

enterprise.Swingle �956! conducted further

experiments on the feasibility of com-rncrcial producing channel catfish usinghigh stocking rates; feeding an artificialfeed composed of soybean oil meal,peanut oil meal, fish meal and distillersdried solublcs; and fertililzing with aninorganic fertilizer. Maximum produc-tion achieved with this method was

1,242 pounds pcr acre. Swingle �959!,Prather �9*1! and others continued re-search on production technology, includ-ing feed formulations, needed for a suc-cessful comniercial catfish farming en-terprise. Although these early diets were

Catfish harvest from a conrrneraal pond in the mississippi deita.

Wellborn � Development of lndNstry 27

not nutritionally complete, they gave satisfactoryresults at the low stocking densities �,000 to 2,000fish per acre! then used.

Education

Because aquaculture is a relatively new farmingenterprise in the United States and requires totallydifferent managerial skills from those normally ac-quired in the production of domestic livestock androw crops, cxtcnsion education programs and assis-tance are necessary for training people on how toproduce, process and market fish. Applied researchis also necessary to develop the information ncixiixito improve production, processing and marketingtechnology.

Early assistance to catfish farmers was providedby Auburn University and the Arkansas Game andFish Commission. But Meyer ef al. �967! pointed outthat "the high number of novices in fish culture hastaxed assisting agencies to the limit and further ex-pansion of the industry can only increase the need fortechnica I assistance."

Thc value of extension aquaculture educationprograms can be illustrated by using those devel-oped in Mississippi for the catfish industry as amodel. Coinmercial production of food-size catFLshstarted in Mississippi in 1965 with the construction ofa 40-acre pond in Sharkey County Wellborn, 1983!.At that time there werc already 9,500 acres of com-mercial catfish ponds in Arkansas Meyer et al�1967!.By 1968 the acres in catfish production werc expand-ing rapidly in the state and farmers went to Missis-sippi State University, the Land Grant University,and requested assistance. As a result of this request,Mississippi State University hired its first extensionfisheries specialist and fisheries researcher. Exten-sion education programs for catfish farmers in Mis-sissippi started in 1969 with a wa ter quality rnanage-rnent workshop. Since tha t time numerous ex tensioncducahon programs on all phases of catfish farming,including processing and marketing, have been con-ducted by specialists, agents and home economists ofthe Mississippi Cooperative Extension Service. Ex-tension programs were also conducted for personnelof lending institutions to educate them as to what wasinvolved in the commercial production of catfish.Additionally, the Mississippi Cooperative ExtensionService Fish Discase Diagnostic Laboratory providedabsolutely essential educational programs and assis-tance to fish farmers on disease diagnosis, treatmentrecommendations and water quality assessment.

Thc role of the extension worker is to provideeducational opportuni ties to citizens of the sta te andassist them in solving problems to enhance the qual-ity of their life, The extension worker also serves asthe liaison bctwccn clientele and researchers at theuniversity. They translate applied research inforrna-tion into layman's terms and take it to their clientele,Extension workers identify problein areas on which

the researcher should focus. Mississippi State Uni-versity has used the mul tidisciplinary a pproach andhas involved all of the appropriate subject matterdepartinents, both in the Mississippi CooperativeExtension Service and the Mississippi Agriculturaland Forestry Experiment Station, in assisting thecatfish farming industry in the state.

The research by many different scientists Cruz,1975; Lovcil, 1971; Prather and Lovell, 1972; Pratherand Lovell, 1973; Tiemcier et aI., 'l 965; Wilson, 1973!,on thc nutritional requireinents of channel catfishwas invaluable since it allowed thc formulation of an

economical, nutritionally complete feed needed forintensive production. Extension education programsand applied research certainly played a major role inthe development of the catfish industry in Missis-slPPl.

MarketingInitially the food-size catfish produced were sold

to local live markets. As acreage increased in the late1960s these local markets were soon saturated andfarmers had to look for other outlets for their fish.Because of the pressure to sell their catfish, somefarmers started small-scale processing on their farmsand began to sell dressed catfish to local restaurantsand grocery stores. In the late 1960s several largecompanies became interested in the processing andma rkcting of farm-raised catifsh as a means of diver-sification. Many of thcseinitial small processing plantsfailed for a variety of reasons.

With the advent of these small processing plantscame efforts to expand beyond local markets to re-gional and national markets. Thcsc early efforts metwi th I irni ted success because of the production meth-ods used. Basically, large fingcrlings were stocked inthe late winter or early spring, fed during the spring,surnmcr and early fall, and then harvested. Thus,catfish were available for sale only during part of theyear. This production method proved to be a majorhandicap in developing markets if catfish farmingwas to become an industry and expand beyond somelocal and regional markets. The growth of catfishprocessing is shown in Figurc 1.

Year-r oun d Prod uction

The sporadic supply of catfish, lack of processingcapacity and suitable market outlets plagued catfishfarmers until about 1974 Wellborn, 1983; Waldrop,1986!, Two things occurred then that had a majorimpact on catfish farming in Mississippi. First, PaulSmith of Yazoo City, Miss., initiated an innovahvcproduction method enabling farmers to supply food-size fish throughout the year; and, secondly, someMississippi catfish farmers formed a cooperative tobuild and operate a feed mill that produced a catfishfeed of a consistently uniform, high quality. Becauseof these two factors, Mississippi catfish farmers claim"industry status" starting in 1974.

Thc development of the multiple harvest or top-

Red Drum Aq<iaculture

350

300

250

0 2000

0 ~ ol50l00

30 69 70 71 72 73 74 75 76 77 78 79 80 Hl 82 b3 84 85 86 87 88 89Years

Figure 1. Millions of pounds of live farm-raised catfish processed hu catfish prnce.".i ng plant s fr<»<i 1969 through 1989 US Krt Catfish, ' ropReporting Hoard, Economicand Statistics Service!

100

60

0 C40

0 2� May Mar. Dec. Dec. Dc~. Dec. Dec. Doc Du<.. July Mar. Apr.1977 1979 1980 1981 I 982 19HH I <!H4 19H, I <!Hr 1'iH7 19HH 1989

Figure 2. Water acres in production of farm-raised catfish in /Vl<ssissipp< from A1a<i 197 to Ay ril 1989.

Welll>orn � Development of Industry 29

ping system allowed farmers to provide processingplants with catfish year-round. Thus, processingplants werc able to expand existing markets and,more importantly, develop ncw markets that re-quired fish all year.

Building of the catfish feed mill by thc farmercooperative allowed catfish farmers to ensure thequality of feed needed for best growth and feedconversions at reasonable prices. Feed was forrnu-lated according to the best knowledge available andwas changed only when applied research has dem-onstrated the value of changing it,

In d us try In Eras tructureCritical to thc development of catfish farming as

a full-fledged industry in Mississippi was the bring-ing together of all of the component parts, i.e., naturalresources, production, extension and applied researchprograms, feed mills, processing capability, cquip-rnent and chemical suppliers, and marketing, in oncgeographic area. This needed infrastructure devel-oped in Mississippi as the result of the close workingrelationship between innovative farrncrs, businessmen, Mississippi Cooperative Extension Service,Mississippi State University, and appropriate stateand federal agcncis.

The growth of the farm-raised catfish industry isevidenced by the increase of the pounds of farm-raised catfish processed annually since 1969 Figure1! and thc increase in water acres in Mississippi in theproduction of farm raised catfish Figure 2!. Manydifferent factors, both overt and subtle, influence thesuccess of any new enterprise. Failure to keep inmind that the ultimate goal of a farmer, businessman, or cnh'cpeneur is making a profit is one of thegreatest impediments to the dcvclopment of a newenterprise. From what I know, it appears that redfishculture as a commercial enterprise is in stage two ofdevelopment. Redfish can be spawned with a mini-mum of trouble, although I do nol know if cost ofproduction has been determined. Suitable and eco-nornical feeds for the production of fingerlings andfood-size fish are still being developed.

It is apparent from the information available inthe manual and from the strong at tendance at thc RedDrum AquacultureConference, that progressisbeingmade in developing extension education and appliedresearch programs, markets for the product, andproduction methods. I feel that the flcdgling red fishinsdustry can certainly benefit from looking closelyat the farm-raised catfish industry to learn where ithad successes and failures.

Literature Cited

Busch, R.L. 1985. Channel catfish culture in ponds. lnC.E. Tucker, Ed. Channel Catfish Culture, ElsevierSciencePublishing Co., inc. N.Y., N.Y. 66 pp.

Ciapp, A. 1929, Some experiments in rearing channelcatfish, Trans, Amer. Fish Soc. 59.114-117.

Crux, F..M. 1975. Determination of nutrient digcstibil-it! in various classes of natural and purified fcM materialsfor channel «atfish, Ph.D. Dissertation, Auburn Univ. 82pp.

Foster, 1'.H. and J,E. Waldrop, '1972. Cost-sire relation-ships in th>; productionof pond-raised catfish for Food. MsAgricuit. & Forest. Exp. Station Bull. 792, 69 pp.

Lcary, 1.1.. 1910. Propagation of crappie and catfish.Trans, Amer. Fish Soc. 39 �9091: pp. 143-148.

Lena, Gerhard. 1947. Propagation of' Catfish. Prog.Fish-Cult 914!:231- 233.

L»vclk R.T. 1971. Calcium and phosph<>us require-ments r>F channel catfish. Fisheries Ann. Rep. Ala. Agr,Fxp, Sta., Auburn Univ�37 pp.

Mcyei, F.P., D.l. Gray, W.P, Mathis, J.M. Martin and8 k. Wells 1967.1'roduction and returns From the commer-cial p rod u> t ion of fish in Arkansas during 1966. Proc. 21 stAnnual Conference of the S.E, Assoc. Game and FishComm.: pp. 525-531

Moblcy, Ben E, 1931. The culture of channel catfish Ictalurus pu>ictatus!. Trans, Amer. Fish Soc 61:171-1 73.

Morris, A.C.1939.Propagationofchanncl«attish. Prog.Fish-Cult., No. 44:23-27.

Prather, E,E, 1961. A comparison of production ofalbino and normal channel catfish. Proc, 15th Ann. Conf.S.E. Assoc. Game & Fish Comm., pp. 302-303.

Vrather, E.E. and R.T. Lovell, 1972, Effcs.-t of vitamintortification in Auburn No. 2 fish Fecxl. Proc. S. E. Assoc.Game and Fish Comm. 25:479483.

Vrather, E.E. and R,T, Lovell. 1973. Response of inten-sively fcd channel catfish tr> diets containing various pra-te>n~nergy ratios. Proc. S. E. Game and Fish Comm, 27:455459.

Swingle, H.S. 1954. Experiments on commercial fishproduction in ponds.8th Ann, Conf. S.E. Assoc. Game andFish Comm.: pp, 69-74.

Swingle, H.S. 1956. Preliminary r csults on the commer-cia1 production o F c'hanne1 catfi sh in pond s. Proc. 10th Ann.Conf, S.E. Game & Fish Comm.: pp. 160-162.

Swingle, H.S. 1959. Experiments on growing tingcrlingchannel catfish to marketable size in ponds. Proc. 12 Ann,Conf. S.E. Game & Fish Comm.: pp, 63-72.

Tiemeicr, O,W., C.W. Deyoe and S Wcardon. 1965,EFFcwts on growth of fingerling channel catfish oF dietscontaining twocnergy and two protein levels. Trans. KansasAcad. Sci. 68:180-186.

Waldrx>p, J.E. 1986. An overview of thc.' farm-raisedcatfish industry in the Unitecl States-an agricultural econo-mists view. In Bilio, M., H, Rosenthai and C.J. Sindcrman, Eds.!. Realism inAquacu Jture: Achievement, constraints,perspectives. Euporean Aquaculture Soc. pp. 519-533.

Waldro p, J.E. and R D,Smith. 1980. An economic analy-sis of producing pond-raised catfish for Foc>d in Missis-sippi: A January1980 update. Dept, of Agricult, Econ. Res.Rcp. MAFES! 103.

Wellborn, T.L., Jr. '1983. The catfish story: farmers, stateservices create new industry. 1983 Yearbook of Agricul-ture, USDA, pp. 298- 305.

Wellborn, T,L�R.M. Durborow, M.D. Crosby and P.W,I'aylor. 1987. Status of fish Farming in Mississippi-Decem-ber 1986. For Fish Farmers, MS Cooperative ExtensionService, 7 pp. mimeo.

Wilson, R V. 1973, Absence of ascorbic acid syiithcsis in«hannel catfish, Ictaluruspunct>rtus, and blue channel catfish,Ictal urus fu rcr>tus. Comp, Biochem, Physiol. 46B: 635-638.Production in thc United State~.