Fish and Wildlife Service

U.S. Department of the Interior

Coastal Ecology GroupWaterways Experiment Station

U.S. Army Corps of Engineers

jA-y Fw$MO\(Q$~LC e..

Biological Report 82(11.108)TR EL-82-4August 1989

Species Profiles: Life Histories and EnvironmentalRequirements of Coastal Fishes and Invertebrates (Mid-Atlantic)

ATLANTIC MENHADEN

by

S. Gordon Rogersand

Michael J. Van Den AvyleGeorgia Cooperative Fishery Research Unit

School of Forest ResourcesUniversity of Georgia

Athens, GA 30602

Project OfficerDavid Moran

National Wetlands Research CenterU.S. Fish and Wildlife Service

1010 Gause BoulevardSlidell, LA 70458

Performed forCoastal Ecology Group

Waterways Experiment StationU.S. Army Corps of Engineers

Vicksburg, MS 39180

and

U.S. Department of the InteriorFish and Wildlife ServiceResearch and Development

National Wetlands Research CenterWashington, DC 20240

This series may be referenced as follows:

U.S. Fish and Wildlife Service. 1983-19 . Species profiles: life historiesand environmental requirements of coastal fishes and invertebrates. U.S. FishWildl. Serv. Biol. Rep. 82(11). U.S. Army Corps of Engineers TR EL-82-4.

This profile may be cited as follows:

Rogers, S. G., and M. J. Van Den Avyle. 1989. Species profiles: life historiesand environmental requirements of coastal fishes and invertebrates (Mid-Atlantic)--Atlantic menhaden. U.S. Fish Wildl. Serv. Biol. Rep.82(11.108).U.S. Army Corps of Engineers TR EL-82-4. 23 pp.

PREFACE

This species profile is one of a series on coastal aquatic organisms,principally fish, of sport, commercial, or ecological importance. The profilesare designed to provide coastal managers, engineers, and biologists with a briefcomprehensive sketch of the biological characteristics and environmentalrequirements of the species and to describe how populations of the species may beexpected to react to environmental changes caused by coastal development. Eachprofile has sections on taxonomy, life history, ecological role, environmentalrequirements, and economic importance, if applicable. A three-ring binder isused for this series so that new profiles can be added as they are prepared.This project is jointly planned and financed by the U.S. Army Corps of Engineersand the U.S. Fish and Wildlife Service.

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

Information Transfer SpecialistNational Wetlands Research CenterU.S. Fish and Wildlife ServiceNASA-Slidell Computer Complex1010 Gause BoulevardSlidell, LA 70458

or

U.S. Army Engineer Waterways Experiment StationAttention: WESER-CPost Office Box 631Vicksburg, MS 39180

iii

CONVERSION TABLE

Multiply

mill-imeters (mm)centimeters (cm)meters (iM)meters (m)kilometers (km)kilometers (km)

square meters (m2

)

square kilometers (km 2 )hectares (ha)

liters (1)cubic meters (m3 )cubic meters (m3 )

milligrams (mg)grams (g)kilograms (kg)metric tons (t)metric tons (t)

kilocalories (kcal)Celsius degrees ('C)

inchesinchesfeet (ft)fathomsstatute miles (mi)nautical miles (nmi)

Metric to U.S. Customary

0.039370.39373.2810.54680.62140.5396

10.760.38612.471

0.264235.310.0008110

0.000035270.035272.205

2205.01.1023.968

1.8( 0 C) + 32

U.S. Customary to Metric

25.402.540.30481.8291.6091.852

To Obtain

inchesinchesfeetfathomsstatute milesnautical miles

square feetsquare milesacres

gallonscubic feetacre-feet

ouncesouncespoundspoundsshort tons

British thermal unitsFahrenheit degrees

millimeterscentimetersmetersmeterskilometerskilometers

square feet (ft 2 )square miles (mi 2 )acres

gallons (gal)cubic feet (ft 3 )acre-feet

ounces (oz)ounces (oz)pounds (lb)pounds (lb)short tons (ton)

British thermal units (Btu)Fahrenheit degrees (*F)

0.09292.5900.4047

3.7a50.02831

1233.0

28350.028.350.45360.000450.9072

0.25200. 5556 (-F

square meterssquare kilometershectares

literscubic meterscubic meters

milligramsgramskilogramsmetric tonsmetric tons

kilocalories32) Celsius degrees

iv

CONTENTS

Page

PREFACE .................................................... iiiCONVERSION TABLE .................. ......................... IVACKNOWLEDGMENTS .......................................................... vi

NOMENCLATURE/TAXONOMY/RANGE ............................................. 1MORPHOLOGY/IDENTIFICATION AIDS ...................................... 1REASON FOR INCLUSION IN SERIES ........................................... 3LIFE HISTORY... ......... ....................... 3

Adult Migration and Spawning ........................................... 3Fecundity .............................................................. 4Eggs and Larvae ......................................................... 4Juveniles and Adults................................................... 6

GROWTH CHARACTERISTICS ................................................... 7THE FISHERY ....... ...................................................... 9

History ................................................................ 9The Catch ......................................... ..................... 9Management ............................................ ................. 10Subpopulations ......................................................... 11

ECOLOGICAL ROLE .......................................................... 12ENVIRONMENTAL REQUIREMENTS ............................................... 13

Temperature, Salinity, and Dissolved Oxygen ............................ 13Substrate and System Features .......................................... 14Environmental Contaminants ............................................. 15Other Factors .............................. ............................ 15

LITERATURE CITED ......................................................... 17

V

)

ACKNOWLEDGMENTS

Technical reviews of this Species Profile were provided by Dean Ahrenholz,Sheryan Epperly, William Hettler, Pernell Lewis, Robert Lewis, and Joseph Smithof the National Marine Fisheries Service, Beaufort, North Carolina, and by WalterNelson, National Marine Fisheries Service, Miami, Florida.

vi

Figure 1. Atlantic menhaden.

ATLANTIC MENHADEN

NOMENCLATURE/TAXONOMY/RANGE

Scientific name: Brevoortia tyrannus(Latrobe)

Preferred common name ......... Atlanticmenhaden (Robins et al. 1980;Figure 1)

Other common names: pogy, moss-bunker, bunker, fat-back, shad,bug-mouth

Class ..................... OsteichthyesOrder ..................... ClupeiformesFamily ............ Clupeidae (herrings)

Geographic range: Temperate coastalwaters from Nova Scotia southward,to Jupiter Inlet, Florida (Dahlberg1970). Atlantic menhaden are sea-sonally abundant in the Mid-Atlantic

* Region (Figure 2). Concentrationsof age 0 fish occur in inshoreestuarine waters along the entireAtlantic seaboard.

MORPHOLOGY/IDENTIFICATION AIDS

Branched dorsal rays, 13-18;branched anal rays, 16-22; pectoralrays, 14-18; pelvic rays, 7; gill fil-aments, 51-66; lateral line scales,40-50; ventral scutes, 29-34; verte-brae, 44-49. Body oblong and com-pressed with a thin belly wall; scaleslarge, coarse, with long slenderpectinations, strongly overlapping andin regular rows; predorsa! scales oneither side of median line enlarged;prominent radiating opercular stria-tions; and pelvic fin rounded withinnermost and outermost rays aboutequal in length (Hildebrand 1963;Dahlberg 1970, 1975).

Color in life: green, brown, orblue-gray, darker on dorsal surface.A dark humeral spot may be followedposteriorly by a series of smaller

1

NEW YORK 0

ATLANTIC OCEAN

0

MILES0 50 100

0 so 100KILOMETERS

M Coastal distribution

HATTERAS

Figure 2. Distribution of the Atlantic menhaden in the Middle Atlantic Region,eastern United States.

2

spots which can fade readily uponcapture. Brevoortia tyrannus can bedistinguished from B. smithi (yellow-fin menhaden, the only other NorthAmerican coast species) because B.tyrannus has a frontal groove, largerand coarser scales in regular rows(therefore, lower scale-relatedcounts), pointed (versus rounded)scale pectinations, a row of lateralspots behind the humeral spot, moregill filaments on the ceratobranchialarch, rounded pelvic fins, and opercu-lar striations. The caudal fin of B.smithi is bright yellow, whereas thecaudal fin of B. tyrannus is not.Fresh B. tyrannus may have a darkeranal fin and more body mucus than doesB. smithi. Atlantic menhaden can bedistinguished from F1 hybrid Atlanticmenhaden (Brevoortia smithi x B.tyrannus) by its longer frontalgroove, lateral spots (absent to fewin hybrid), and a rounded ventral fin.Dahlberg (1970, 1975) provided addi-tional measurements and descriptionsof qualitative characters. Jones etal. (1978) gave detailed descriptionsof Atlantic menhaden developmentalstages (egg through adult). Hettler(1984) gave meristic and morphometricdescriptions of Atlantic menhadenlarvae and juveniles.

REASON FOR INCLUSION IN SERIES

Atlantic menhaden constituteabout 25% to 40% of the combinedannual landings of Atlantic coast andGulf of Mexico menhaden species, whichcollectively comprised the largestcommercial fishery by weight andeighth largest in dollar value in1984 in the United States (NMFS1985). They are important prey formany other fish species and areseasonally important components ofestuarine and shelf fish assemblages.Atlantic menhaden depend on habitatsalong the entire eastern seaboard andadjacent shelf waters throughouttheir life cycle and use estuaries asnursery areas. Some spawning occurs

in estuarine zones and nearshore shelfwaters northward from Chesapeake Bay.Due to the species' great abundance,extensive migration patterns, andimportance as a prey species, theAtlantic menhaden influences theconversion and exchange of energy andorganic matter within biologicalsystems throughout its range (Petersand Schaaf 1981; Lewis and Peters1984).

LIFE HISTORY

Terminology used to describe lifehistory stages conforms to that usedby Lewis et al. (1972) and Moyle andCech (1982).

Adult Miqration and Spawning

Knowledge of timing and locationof spawning has been mainly obtainedfrom collections of adult females thatwere spent or contained maturing ova(Higham-and Nicholson 1964; June 1965;Dahlberg 1970) as well as from collec-tions of eggs and larvae (Reintjes1961, 1968; Herman 1963; Kendall andReintjes 1975; Ferraro 1980a,b; Judyand Lewis 1983). Data on movement andpopulation age or size structure havebeen obtained from distribution ofpurse-seining effort (Roithmayr 1963);frequencies of lengths, weights, andages in catches (June and Reintjes1959;. June 1972; Nicholson 1971,1972); and returns from extensivetagging experiments (Dryfoos et al.1973; Kroger and Guthrie 1973;Nicholson 1978). Atlantic menhadenundergo extensive north-south seasonalmigrations and inshore-offshoremovements along the Atlantic seaboard.Schools are composed of fish ofsimilar size and age. Migrationpatterns are also related to spawninghabits, and some spawning occurs everymonth of the year.

Atlantic menhaden of allcongregate off North Carolina

agesfrom

3

November to March and some spawn therein shelf waters that are 100 to 200 mdeep, probably within 70 m of thesurface (Reintjes and Pacheco 1966).All eggs of Atlantic menhadencollected by Judy and Lewis (1983)were taken at depths of 10 m or less.The spawning is heaviest off CapeLookout, North Carolina, from Decemberthrough February. Adults then moveinshore and northward in spring andstratify by age and size along theAtlantic seaboard. Adult menhadenhave been collected from estuaries,and some move as far inland as thebrackish-freshwater boundary. Theoldest and largest fish migrate thefarthest, some moving as far north asthe Gulf of Maine.. Adults that remainin the South Atlantic Region movesouthward and reach northern Floridaby fall. Representatives of all ageclasses return to the shelf waters ofthe South Atlantic Region in latefall.

During migration northward inspring, spawning occurs progressivelycloser inshore; by late spring, somefish spawn within coastal embaymentsof the North Atlantic Region. Thereare definite spring and fall spawningpeaks in the Middle and North AtlanticRegions, and some spawning occursduring the winter in the shelf watersof the Mid-Atlantic Region. Temporaland spatial segregation of spawningactivity provides a mechanism for theexistence of races (= subpopulations).Higham and Nicholson (1964) and Schaaf(1979) have speculated that a femalemay spawn more than once in a season.

Fecundity

Higham and Nicholson (1964)reported values of 38,000 to 631,000ova per fish, and June (1961a) gavevalues of 40,000 to 700,000 ova perfish; estimates depend on the size ofthe fish. Dietrich (1979) reportedfecundities of 116,000 to 568,000ova per fish at age I to age V,respectively (Table 1). Higham and

Table 1. Estimated fecundity ofAtlantic menhaden at different ages(from Dietrich 1979).

Number of eggs per female(thousands)

Age Mean Range N

1 115.8. 26.5 - 250.7 21

II 177.4 39.2 - 368.8 34

1I1 302.8 127.7 - 458.3 33

IV 308.6 142.7 - 514.0 12

V 568.4 --- 1

Nicholson (1964) gave the followingequation for the estimation of fecun-dity (F = ova per fish; FL = forklength, mm):

In F = 7.2227 + 0.0176 FL

r2 = 0.726.

Dietrichequationsweight ofFL = fork

(1979) gave the following(W = wet body weight less

ovaries, g; A = age, years;length, mm):

F = 488 Wr2 = 0.916

F = 92,592 Ar 2 = 0.879

In F = 8.6463 + 0.0120 FL

r2 = 0.871.

Eggs and Larvae

Eggs of the Atlantic menhaden arepelagic and have been reported tohatch in 2 days at unspecified temper-atures (Kuntz and Radcliffe 1917), 2.9days at 15.5 'C (Ferraro 1980a), andat 2.5 to 2.9 days at an averagetemperature of 15.5 CC (Hettler1981).

4

Survival of laboratory rearedAtlantic menhaden embryos to hatchingis very low (2% to 45%); 49% to 94% ofmortality occurs before blastoporeclosure (Ferraro 1980a).

Atlantic menhaden larvae beginfeeding on individual zooplankters(Reintjes and Pacheco 1966) about 4days after hatching when the yolk sacis almost absorbed, the eyes arepigmented, and the mouth is func-tional (Hettler 1981). Larvae in theSouth Atlantic Region enter estuariesafter 1 to 3 months at sea (Reinties1961) at fork lengths (FL) of 14 to34 mm (Reintjes and Pacheco 1966);fish longer than 30 mm FL are thenalready metamorphosing to the pre-juvenile morphology (Lewis et al.1972). This migration into estuariesoccurs from May through October inthe North Atlantic Region, Octoberthrough June in the Mid-Atlantic, andDecember through May in the SouthAtlantic Region (Reintjes and Pacheco1966). As they grow, the larvaeprobably feed on progressively largerzooplankters (Kjelson et al. 1975).

Young fish move into the shallowportions of estuaries including rivershoals and the heads of small tidalcreeks (Massmann 1954; Massmann et al.1954; June and Chamberlin 1959;Pacheco and Grant 1965; Wilkens andLewis 1971; Lewis et al. 1972;Weinstein 1979; Weinstein et al.1980; Rozas and Hackney 1984; Rogerset al. 1984). They apparently preferSpartina, Juncus, and the vegetationtypical of fresh tidal marshes andswamps (Taxodium, Typha, Nyssa)Peltandra, Rumex, Sagittaria, Zizania)over vegetated habitats in open water(Adams 1976; Weinstein and Brooks1983). While in estuaries, Atlanticmenhaden grow and metamorphose througha prejuvenile stage into juveniles.

Several studies have reportedabundances of young menhaden that werehigher in portions of estuaries withthe lowest salinity <5 ppt (Lewis etal. 1972; Weinstein 1979; Weinstein et

al. 1980). Massmann et al. (1954)reported that the abundance of pre-juveniles was higher above than belowthe brackish-freshwater boundary, andRogers et al. (1984) showed that thispattern persists during high riverdischarge. Massmann et al. (1954),Rozas and Hackney (1984), and Rogerset al. (1984) provided evidence thatprejuvenile Atlantic menhaden selecttidewater areas that are fresh or oflow salinity. Only fish of prejuve-nile lengths were resident in lowsalinity river shoals and in inter-tidal creeks (Lewis et al. 1972).This phenomenon persisted for about 4months (Rogers et al. 1984).

A "critical period" of survivalin young fishes was first defined byHjort (1914) and discussed for clupei-form fishes by Schumann (1965) andMay (1974). Menhaden, like mostfishes with high fecundity and littleparental care, hatch in an undevelopedstate. Such fish typically rely onenergy contained in the yolk-sac for4-5 days after hatching, after whichthey are sufficiently developed tomore efficiently feed on an externalfood supply (Schumann 1965). Feedingof the youngest Clupea, Engraulis,and Sardinops larvae depends largelyon food availability; fish will eatto capacity in the presence of highfood concentrations and starve inthe absence. of high concentrationsbecause they are unable to move aboutto search for food. A routine searchpattern is initiated only after anencounter with or capture of a foodparticle. Given the heterogeneity indistribution of pelagic plankters andthe inability of many clupeiformfishes to cope with low food concen-trations, menhaden probably have acritical period of larval survival.Year-class strength may be partlydetermined during this period. Thisproblem is most likely to occur whenlarvae are spawned offshore or sweptoffshore after having been spawnednearshore. Individual larval condi-tion factors (weight-length ratios)increase rapidly when the fish enter

5

K

an estuary (Lewis and Mann 1971).Metamorphosis is not totally dependenton low salinity; Hettler (1981) rearedAtlantic menhaden from eggs tojuveniles using water with a highsalinity. However, metamorphosisrarely is successful outside thefood-rich, low-salinity estuarinezones (Kroger et al. 1974).

No data are available that linksurvival at yolk sac absorption toyear-class strength or that enablequantitative estimates of mortalityfrom predation or starvation (May1974). Minimum food concentration forinception of feeding activity is notknown, and survival curves do notexist for larval Atlantic menhaden.Nelson et al. (1977), however,developed an environment-recruit modelin an attempt to explain variation inyear-class strength. Since larvalmenhaden are thought to depend largelyon wind-driven (Ekman) transport toreach estuarine nursery grounds(particularly in the Middle and SouthAtlantic Regions), the modelincorpo-rated four variables: (1) the knownspawning times and locations; (2) windvectors specific for year, time, andlocation; (3) year- and time-specificdischarges of major tributary systems;and (4) the minimum sea surfacetemperature at the mouth of DelawareBay. A survival index was calculatedas the ratio of observed recruitmentto the fishery (age I) to that predic-ted by a Ricker (1954) spawner-recruit(density-dependent) model. The magni-tude of the index "should reflectthose environmental (density indepen-dent) effects which influence survivalof menhaden from the time of spawningto the time of recruitment to thefishery at age I" (Nelson et al.1977). The model explained 84% of thevariation in the survival index forthe years investigated (1961-71);Ekman transport was the principalcomponent. The correlation has notbeen statistically significant inrecent years (D. Vaughan, NationalMarine Fisheries Service (NMFS),

Beaufort, Northcomm.).

Carol i.na ; pers.

Survival of Atlantic menhaden toage I has been estimated by comparingestimates of population size of age Ifish (based on a virtual populationanalysis that incorporated data fromcommercial landings) with number ofeggs predicted to have been spawnedin previous years. Estimates ofrecruits per million eggs spawnedhave ranged from 27 to 159 (Nelson etal . 1977) and from 78 to 282(Dietrich 1979).

Juveniles and Adults

Metamorphosis marks a change infeeding mode from capturing individualzooplankton to filter-feeding (Juneand Carlson 1971; Durbin and Durbin1975). This shift is accompanied by aloss of teeth,, an increase in thenumber and complexity of gill rakers,and an increase in the complexity andmusculature of the digestive tract(June and Carlson 1971). Prejuvenilesare somewhat intermediate in feedingmode (June and Carlson 1971) and bodystructure (June and Carlson 1971;Lewis et al. 1972).

Juveniles begin congregating indense schools as they leave shoalareas. Most emigrate from estuariesfrom August through November (earliestin the North Atlantic Region) atlengths of 55 to 150 mm FL (June1961a) or 55 to 140 mm total length(TL) (Nicholson 1978). Nicholsonstated that most emigrants are 75 to110 mm TL. As judged by the resultsof extensive tagging, many age 0 fishmigrate southward along the NorthCarolina coast in late fall andearly winter (Nicholson 1978). Fishin the southernmost portion of theSouth Atlantic Region, however,showed less offshore migration(Dahlberg 1970), and tagging resultsindicated that juveniles leaving theestuaries of the South Atlantic Regionand the North Atlantic Region may not

6

move far north or south during theirfirst year (Nicholson 1978). Larvaeentering estuaries late in the seasonmay remain in the estuary one addi-tional year and emigrate at. age I.Some juveniles and adults are found insounds and bays along the SouthAtlantic Region during mild winters(June 1961a). Fish leaving estuariesalong the entire Atlantic seaboardeventually disperse throughout most ofthe geographic range (Nicholson 1978).

Most Atlantic menhaden reachmaturity by the end of their secondfull year. About 10% were found tobe capable of spawning at age I (latein the year), and 90% at age II(Higham and Nicholson 1964). Fish ofall ages, however, are found in themigrating schools. Although Atlanticmenhaden can live 8 to 10 years (Juneand Roithmayr 1960), fish older thanage IV had been rare in the commercialcatch. As stocks rebuild, however,age V and VI fish are becoming morecommon and may be locally abundant inthe North Atlantic and North Carolinafall fishery (Powers 1984).

Adult feeding behavior isaffected by food availability (Durbinand Durbin 1975; Durbin et al. 1981).Swimming speed is increased at higherfood concentrations, and associatedenergetic costs rise exponentially.Modeling studies have suggested thatAtlantic menhaden maximize growthrate, not efficiency (Durbin andDurbin 1983), and that efficiency ofdietary assimilation changes seasonal-ly with the quality of available food.The Atlantic menhaden has behavioral,physiological, and morphologicaladaptations for an active migratoryexistence in waters with pronouncedseasonal and spatial variation in foodabundance. It has large lipidreserves that are seasonally assimi-lated (Durbin and Durbin 1983), a bodyshape adapted for continuous swimming,and copious body mucus (Dahlberg1970).

GROWTH CHARACTERISTICS

Growth rates vary among years andlocalities throughout the species'range (June and Reintjes 1959, 1960;June 1961b; June and Nicholson 1964;Nicholson and Higham 1964a, 1964b,1965; Nicholson 1975; Reish et al.1985). The age of Atlantic menhadencan be determined from annual scalemarkings (McHugh et al. 1.959; June andRoithmayr 1960; Kroger et al. 1974).

Fish of the same age are progres-sively larger in more northerly fish-eries (Nicholson 1978; Reish et al.1985), but mature at smaller sizes inmore southerly areas. Minimum size atmaturity was 180 mm FL in the SouthAtlantic and 210 mm FL in the MiddleAtlantic (Higham and Nicholson1964). There is evidence that growthrates have changed in response tofishing pressure: fish of the same agewere larger in the late 1960's andearly 1970's than in the late 1950'sand early 1960's (Nicholson 1975).Reish et al. (1985) indicated thatgrowth rates do not depend upon fishabundance. Atlantic menhaden in yearsof high abundance probably are smallerthan Atlantic menhaden in years of lowabundance because the former weresmaller at the time of recruitment,not because of any difference ingrowth rates after recruitment.

Growth has been shown to be allo-metric in larval, prejuvenile (Lewiset al. 1972), juvenile (Lewis et al.1972; Epperly 1981), and adult stages.Reish et al. (1985), however, reportedthat Atlantic menhaden exhibitedgrowth that was close to isometric,although growth appeared to becomeallometric with increasing age. Threedifferent "stanzas" of growth in youngmenhaden were reported by Lewis et al.(1972), with inflection pointsat 30 and 38 mm TL (70 and 469 mgweight). These points served as thebasis for their division of the lifehistory stages. Balon (1984) pointedout that these are functional, not

7

arbitrary, divisions. Lewis et al.(1972) cited an unpublished manuscriptthat stated that the relationshipbetween length andweight is similarfor juveniles and adults. Length-weight conversions can be made usingthe appropriate equation in Table 2.

Atlantic menhaden growth beginsin spring and ends in fall, as thewater temperature crosses an approxi-mate 15 *C threshold (Kroger et al.1974). At Beaufort Inlet, North Caro-lina, age 0 fish ranged from 40 to 185

mm TL at the end of the growing sea-son, depending on when they were

Table 2. Weight-length regressions for(log e length).

spawned and entered the estuary.Young of the next year class arrivedin the spring only 20 to 30 mm TLshorter than the smallest fish of theprevious year class. These factors,combined with differences in larvalgrowth rates (Lewis et al. 1972; Kro-ger et al. 1974) and latitudinal dif-ferences in growing season, probablyexplain the observed differences insizes of fish of the same "age" withina single year's catch. See Durbinand Durbin (1983) and the section onLife History for discussions ofgrowth in relation to feeding,environment, and body morphology.

Atlantic menhaden; loge weight = a + b

Types ofmeasurement units

Location Weight Length a b(wet) (mm)

White Oak River Estuary, NC'

Larvae (9 30 mm TL) mg TL -8.110 3.605

Prejuveniles (30-38 mm TL) mg TL -16.964 6.308

Juveniles (a 38 mm TL) mg TL -5.230 3.145

Fall and winter spawners and offspring g SL -10.884 3.067(Middle, S. Atlantic Regions) 2

North Atlantic spring spawners g SL -11.240 3.145and offspring2

Middle Atlantic spawners and g SL -11.037 3.103offspring

2

South Atlantic spawners and g SL -10.579 2.995offspring

2

All spawners, for the fishery g SL -12.075 3.215as a whole

3

'Lewis et al. 1972.2 Epperly 1981.3 Douglas Vaughan, National Marine Fisheries Service, pers. comm.

8

Atlantic menhaden reach lengthsof about 500 mm TL and weights of over1,500 g at ages of 8 to 10 years.Cooper (1965) collected an 8-year-oldthat measured 470 mm TL and weighed1,674 g.

THE FISHERY,

History

The Atlantic menhaden fishery wasfirst established in the late 1600'sand early 1700's to obtain fish foragricultural fertilizer (Frye 1978).In the early 1800's, an industry wasdeveloped to obtain oil f 'rommenhaden (Goode 1879; Goode and Clark1887), and by 1869 there were 90reduction plants in North Carolinaalone (June 1961a). Today thisspecies contributes 25% to 40% of thelandings in the largest commercialfishery (by weight, Brevoortiaspecies) in the United States. Annuallandings for 1979 to 1981 averagedabout 400,000 metric tons and $38million in market value (NMFS 1980,1981, 1982, 1983, 1984, 1985). Plantsthat process Atlantic. menhadenproducts currently operate from Maineto Florida. About 96% *to 918% of thecatch is sold to livestock and cos-metic interests as fishmeal, solubleproteins, and oils; the rest is usedin pet food products and as fish bait(NMFS 1980, 1981, 1982, 1983, 1984,1985). Most of the landings are madewith purse seines. Federal efforts tocollect data for management of Atlan-tic menhaden began in 1955 (June1957).

The Catch

The Atlantic menhaden fishery hastwo annual phases: a summer and fallfishery from Maine to northernFlorida, and an intensive fall andwinter fishery off North Carolinabetween Cape Lookout and New RiverInlet (June 1961a; Nicholson 1978).Landings for the entire fishery have

recently included primarily age I andII fish. During the summer fishery inthe South Atlantic Region, fish caughthave been mostly ages I and 11 exceptin 1984 when a large number of age 0fish were caught; the north Floridalandings have been composed mostly ofage I fish. Concurrently,' fish in theChesapeake Bay area and the southernportion of the Mid-Atlantic Region arealso age I and II, although they arelonger and heavier on the average.Some fish of ages III and IV arepresent in an early spring pound-netfishery in Chesapeake Bay. Most ofthe fish caught in the northernportion of the Mid-Atlantic Region areof ages 11 and III, the age II fishbeing larger than those to the south.For the entire Atlantic menhadenfishery, the average percentage ofnumbers of age I and 11 fish between1955 and 1984 was 78.4%; the range was51.4% (in 1961) to 95.9% (in 1970).Numbers of age 0 fish composed 25% ormore of the catch for the entirefishery in 1955, 1966, 1979, 1981,1983, and 1984 (D. Vaughan, NationalMarine Fisheries Service; pers.comm.). -The north Atlantic fisheryoperates from mid-June through Octoberand primarily exploits fish of age IIIor older. The purse seine fisherynorth of Cape Hatteras is over by lateNovember. Age 0 fish begin to bevulnerable to the fishery during latefall and winter from Chesapeake Baysouth. The North Carolina fallfishery is composed of fish of all ageclasses; age 0 fish have predominatedsince 1971, except in 1980 and 1982.

Atlantic menhaden stocks weredrastically reduced during the 1960's.Annual landings dropped from 671,400metric tons in 1955 to about 200,000metric tons per year from 1966 through1969 (Nelson et al. 1977). As thepopulation size decreased, the agestructure also changed. Fish olderthan age III became scarce and fisholder than age IV were practicallynon-existent even in the NorthCarolina fall-winter fishery. Manynorthern processing plants closed

9

down -- especially those in the NewEngland area, where the fisherydepended on fish of ages III and IV(Henry 1971; Nicholson 1975). Age Iand II fish constituted the bulk ofthe landings and age 0 became moreimportant (Nicholson 1975). Thestocks began to recover in the early1970's, when age III fish againappeared in North Atlantic catches.The first significant Maine landings(3,100 metric tons) since the 1960'soccurred in 1973 (NMFS 1973, 1974,1975; Nelson et al. 1977). NorthAtlantic (from Cape Cod, Massachu-setts, to Cape Breton, Nova Scotia)landings in 1929-71 correlatedstrongly with 3-year-lagged localwater temperatures and mixing factors%for St. Lawrence River inputs'(Sutcliffe et al. 1977). These NorthAtlantic fish, though vulnerablethroughout the fishery, may be aunique biological stock. Catchescontinued to improve into the early1980's; however, the size of thereproductive stocks (ages III and IV)remained low (Atlantic MenhadenManagement Board [AMMB] 1981; NMFS1983, 1984, 1985). Heavy exploitationresults in smaller and fewer fish(June 1972) and higher catchabilitycoefficients (Schaaf 1979). Theimplications of these phenomena arenot fully understood; however, a recentmodel showed that pollution stress maygreatly reduce first-year survival rate(Schaaf et al. 1987).

Management

More than 50% of the annuallandings of Atlantic menhaden are. fromwithin State territorial waters,mostly from the Chesapeake Bay area(R. B Chapoton, NMFS, Beaufort, NorthCarolina; pers. comm.). This fact,combined with the migratory nature ofthe species and the dependence ofnorthern fisheries on. escapement ofage I and II fish from fisheries inthe South Atlantic Region andChesapeake Bay (Nicholson 1978), makesregulation a _compromise, situationbetween the industry and Federal and

State agencies; only the states havefinal regula.tory power.

The stocks have generally beenunmanaged. Nelson et al. (1977)suggested that the fishery is somewhatself-requlating in that reducedcatches bring about reduced effort andplant closures, allowing the stocks torecover. They stress that propermanagement practices could reduce thechance of repeating the mistakes ofthe past and prevent a crash in thefishery. In addition, Schaaf (1975)pointed out that "allowing the fisheryto be controlled . . . by the econo-mics of free market competitionassures that (1) the average profitfor the whole industry will be zero• . . and (2) there is no mechanism toprovide for protection of theresource, since if either costs godown or value goes up new effort canafford to enter the fishery andeventually may exceed the BBEP[Biological Break-Even Point, a pointat which the fishery collapses]."Schaaf (1975) also warned that thelevel of effort when the fisherycollapsed in the 1960's might bemaintained during the mid-1970's evenif catches dropped to 200,000 metrictons per year because product priceswere higher at that time. He urgedimplementation of a flexible quotasystem coordinated by the AtlanticStates Marine Fisheries Commission(ASMFC) that would allow the stocks tocontinue to rebuild while effortregulation mechanisms were studied.

The management option endorsed bythe ASMFC, Option 7 of the AtlanticMenhaden Advisory Committee, proposedthe following closing dates to protectage 0 recruits: the week endingbetween October 4 and 10 for the NorthAtlantic, the week ending betweenOctober 11 and 17 for the Mid-Atlantic, the week ending betweenNovember 8 and 14 for Chesapeake Bay,and the week ending between December13 and 19 for the South Atlantic.This closure would primarily affectthe North Carolina late-fall fishery.

10

To be effective, the regulation stillneeds to be implemented in NorthCarolina; it has been implementedalready in six states (Vifrginia andnorth). National Marine FisheriesService personnel 'have analyzed theeffects that Option 7 would have byusing data for 1976-82. Increases inyield-per-recruit ranged from 0.4% to>10% and varied with strength of thetargeted age 0 year class and thetiming of arrival at the NorthCarolina fall fishery. These esti-mates may be low because of samplingerrors, but would be substantiallygreater only in years with veryabundant age 0 cohorts that arrive atthe North Carolina fall fisheryafter the mid-December closing date(D. Vaughan, pers. comm.). Thismanagement strategy is an extension ofNMFS recommendations made since theearly years of study. The continuedexistence of the Atlantic menhadenfishery despite heavy exploitation isquite unlike that of other well-studied clupeid stocks (Schaaf 1979),and probably is a product of differ-ences in density-independent phenomena(e.g., reproductive strategy). Inspite of the difficulties involved inmanaging such a complex resource, theprospects of developing a workablemanagement plan are good, primarilydue to the capability of the stocks torespond to conservation of age O-fish(Schaaf 1975). However, the coopera-tion of affected states is necessaryto effectively implement such a plan.

Annual maximum sustained yield(MSY) from historic catch-effort datahas been estimated by Schaaf andHuntsman (1972) at 600,000 metrictons and by Schaaf (1975) at 560,000metric tons. Estimates incorporatingthe use of a Ricker (1954) spawner-recruit (density-dependent survivor-ship) model were given as 380,000metric tons per year by Schaaf andHuntsman (1972) and averaged 419,000metric tons per year on the basis ofknown survivorship in 1961-71 (Nelsonet al. 1977). The recruit-environmentmodel developed by Nelson et al.

(1977) likewise averaged 419,000metric tons per year (1961-71) and isproposed to offer a way of "fine-tuning" predicted catch on a yearlybasis by constantly updating yieldestimates. The vulnerability of theAtlantic menhaden fishery. to fluctua-tions in year class strength was firstpointed out by June (1961a). It hassince been stressed that the mainte-nance of a healthy stock of spawning-age fish should be a primary concernof management (Schaaf and Huntsman1972; Schaaf 1975; Nelson et al. 1977;Vaughan 1977). Good stocks ofspawning-age fish would bring multiplebenefits, including higher reproduc-tive potential (decreasing the effectsof years with poor recruitment),decreased vulnerability to weak yearclasses, and increased weight oflandings due to a higher contributionof older fish to the catch. Recentcalculations of MSY remain near450,000 metric tons (D. Vaughan,pers. comm.). I

Annual instantaneous natural mor-tality was estimated at 0.36 (1955-64)by Schaaf and Huntsman (1972) and at0.42 for the 1955 year class by Nelsonet al. (1977). Their respective esti-mates of annual instantaneous fishingmortality were 0.82 to 2.14 (1955-64)and 0.36 for the 1955 year class. Acombination of these data yieldstotalinstantaneous mortality estimatesranging from 1.18 to 2.56, whichcorrespond to total annual mortalityrates of 69% to 92% (ages I - VI).Reish et al. (1985) estimated annualnatural mortality at 0.54 for latejuveniles (9-23 months), 0.15 for ageI (12-23 months), 0.49 for age II(24-35 months), and 0.52 for age 3+(36+ months). Except for the age Ifish, these estimates were comparablewith the annual mortality of 0.52obtained from mark-recapture data byDryfoos et al. (1973).

Subpopulations

Because a genetically distinctstock can have its own homogenous

I1

vital parameters of recruitment,growth, and mortality (Cushing 1968),identification of the stock (=subpopu-lations) and stock-specific biologicaltraits is necessary for proper manage-ment. Various authors have proposedthe existence of two to five Atlanticmenhaden. subpopulations on the basisof meristic and morphometric compari-sons (June 1958, 1965; Sutherland1963; Higham and Nicholson 1964; Juneand Nicholson 1964; Dahlberg 1970;Epperly 1981). Dahlberg (1970)reported a distinct subpopulation ofAtlantic menhaden south of CapeCanaveral' in the vicinity of theIndian River, Florida. Nicholson(1978) stated that the extensivenorth-south migrations, latitudinalstratification by age and size in thesummer, and intermingling of all ageclasses south of Cape Hatteras inwinter preclude the existence of morethan one stock. Epperly (1981),however, provided electrophoretic aswell as meristic and morphometric datathat indicated significant differencesbetween fish spawned in the watersnorth of Long Island, New York, duringthe spring and those spawned in thefall and winter in the South andMid-Atlantic Regions. Other groups --fall-spawning fish of the Gulf ofMaine and spring-spawning fish of theMid-Atlantic Region -- may also bedistinct subpopulations, but thisaspect has not been fully investi-gated. Menhaden species hybridizereadily (Turner 1969; Dahlberg 1970),but Atlantic x yellowfin hybrids havebeen recorded only as far north asBeaufort, North Carolina (Dahlberg1970). Apparently, hybrids do notoccur in the Mid-Atlantic Region.

ECOLOGICAL ROLE

Atlantic menhaden occupy twodistinct types of feeding nichesduring their lifetime. They are size-selective plankton feeders as larvaeand filter feeders as juveniles andadults. Data on the food of larvae

before they enter the estuary do notexist. After entering the estuary,Atlantic menhaden larvae appear to beextremely selective for prey of cer-tain sizes and species. Larvae fromthe Newport River Estuary, NorthCarolina, 26 to 31 mm TL (x = 29 mmTL), consumed copepods and copepoditesof only four taxa, which composed 99%by numbers and volume of their gutcontents (Kjelson et al. 1975). Theseprey items, ranging from 300 to 1200.um in length (ý = 750 um), were eatendespite an abundance of copepodnauplii, barnacle larvae, and smalladult copepods in plankton tows.Larvae that were offered copepods inthe laboratory ignored all other fooditems, including Artemia and Balanusnauplii (June and Carlson 1971).Larval menhaden in the Newport RiverEstuary, North Carolina, fed primarilyduring daylight (Kjelson et al. 1975).

Juvenile and adult Atlanticmenhaden strain particulates from thewater column with a complex set ofgill rakers. The rakers can sieveparticles down to 7-9 pm in size(Friedland et al. 1984), includingzooplankton, larger phytoplankton, andchain-forming diatoms. Biochemicalanalyses indicated that the gutcontents of juveniles vary with preyavailability; reliance on zooplanktondecreases as the fish move from openwaters to. marshes (Jeffries 1975).Atlantic menhaden may also be capableof eating epibenthic materials (Edgarand Hoff 1976). Peters and Schaaf(1981) speculated that the annualphytoplankton and phytoplankton-basedproduction in east coast estuaries isnot sufficient to support the juvenileAtlantic menhaden population duringits residency and that the abundantorganic detritus may be eaten. Lewisand Peters (1984) reported thatjuvenile Atlantic menhaden in NorthCarolina salt marshes ate primarilydetritus.

The roles of Atlantic menhaden insystems function and community

12

dynamics have received littleattention. Larvae and juveniles areseasonally important components ofestuarine fish assemblages (Tagatz andDudley 1961; Cain and Dean 1976;Bozeman and Dean 1980). Estimates ofthe mean daily ration for larvae rangefrom 4.9% (Kjelson et al. 1975) to 20%(Peters and Schaaf 1981) of wet bodyweight. Assimilation of ingestedenergy exceeded 80% for plant andanimal material (Durbin and Durbin1981). Because of their tremendousnumbers, individual growth rates, andseasonal movements, these fishannually consume and redistributelarge amounts of energy and materials,including exchanges between estuarineand shelf waters.

Kjelson et al. (1975) noted thatthe copepod taxa preferred by larvalmenhaden and other species decreasedfrom a mean value (2 years) of 81% to48% of the total zooplankton biomassduring the period of larval residence.They speculated that this decrease maybe partly explained by larval feeding.Durbin and Durbin (1975) suggestedthat Atlantic menhaden in coastalwaters may also alter the compositionof plankton assemblages by grazing oncertain size ranges.

Important Atlantic menhadenpredators include bluefish (Pomatomussaltatrix), striped bass (Moronesaxatilis), bluefin tuna (Thunnusthynnus), and sharks (Reintjes andPacheco 1966). Atlantic menhadenoccurred in 13% of stomachs of sandbarsharks (Carcharhinus plumbeus). About46% of the menhaden were consumed wholeand were 5-10 cm TL. Menhaden consumedin more than one piece or partiallyconsumed were 5-17 cm TL. This preywas the second most frequently consumedtype for sandbar sharks (Medved et al.1984). Because Atlantic menhaden areeaten by predators in several eco-systems, they are a direct pelagic linkin the food web between detritus andplankton and top predators.

ENVIRONMENTAL REQUIREMENTS

Temperature, Salinity, and DissolvedOxygen

Atlantic menhaden occur through awide range of physicochemical condi-tions. Several studies have raisedquestions about limits of toleranceand optimum conditions. June andChamberlin (1959) and Reintjes andPacheco (1966) reported that larvalmenhaden did not enter estuarinewaters at temperatures below 3 *C.Many studies have noted an affinity ofyoung menhaden for low salinity waters(see the Life History section).Wilkens and Lewis (1971) speculatedthat larval menhaden require lowsalinity water to metamorphoseproperly, and Lewis (1966) found thatalthough larvae metamorphosed in.salinities of 15 to 40 ppt, one-thirdof the juveniles developed slightlycrooked vertebral columns. However,larvae held in the laboratory at 25 to40 ppt metamorphosed completely withno abnormalities (Reintjes and Pacheco1966; Hettler 1981); and larvaetrapped in a natural cove at Beaufort,North Carolina, transformed intojuveniles at 24 to 36 ppt (Kroger etal. 1974).

Salinity affects temperaturetolerance, activity, metabolism, andgrowth. . Low salinities decreasedsurvival at temperatures below 5 CC,and survival was poor at 6 'C infreshwater (Lewis 1966). The effectof salinity on upper temperaturetolerance was not significant (Lewisand Hettler 1968). Larvae thatHettler (1976) reared at 5 to 10 pptexhibited significantly higher activi-ty levels, metabolic rates, and growthrates than those reared at 28 to 34ppt. Lewis (1966) also noted slowergrowth at high salinities. Subtlephysiological adaptations to lowsalinity may be an evolutionaryresponse to larvae "seeking" thefood-rich estuarine environment.Rogers et al. (1984) noted that pre-juveniles of many fishes, including

13

those of Brevoortia species, enteredestuarine habitats during seasonalpeaks of freshwater influx when thearea of low-salinity and fresh tidalwater was greatest.

An important management consider-ation is that, during the evolution ofthe Atlantic menhaden, estuarine zonesreceived freshwater from contiguouswetlands and riverine systems. How-ever, channelization, diking of rivercourses, ditching and draining ofmarginal wetlands, and urbanizationhave reduced the freshwater retentioncapacities of coastal wetlands.Furthermore, extensive filling ofestuarine marshlands has diminishedthe area receiving runoff in manylocations. In combination, thesechanges cause rapid discharge of highvolumes of freshwater during briefperiods and reduced amounts of fresh-water at other times. High inflows,particularly those that occur in earlyspring after the arrival of pre-juvenile menhaden, can expose fish toextreme fluctuations of temperature,turbidity, and other environmentalconditions. Although the effects ofaltered freshwater flow regimes onAtlantic menhaden are not known,effects on other estuarine-dependent,offshore-spawned fishes range fromdisappearance (Rogers et al. 1984) todeath (Nordlie et al. 1982). Theseeffects are also mediated by tempera-ture (Nordlie 1976).

Salinities of 10 to 30 ppt didnot affect developing embryos, thoughtemperature did (Ferraro 1980a).Mortality of embryos was complete attemperatures less than 7 °C and wassignificantly higher at 10 'C than at15, 20, and 25 °C. Time to hatchingwas significantly shorter at eachprogressively higher temperature.Surface temperature in the spawningareas of the South Atlantic Regionduring the months of highest eggcapture were generally 12 to 20 'C(Walford and Wicklund 1968). Thelowest temperatures at which Atlanticmenhaden eggs and larvae were collec-

ted in the North Atlantic Region werebetween 10 and 13 'C (Ferraro 1980a).The temperature range for the MiddleAtlantic Region was 0 to 25 'C, butmost eggs and larvae were collected at16 to 19 'C (Kendall and Reintjes1975).

The limits of larval temperaturetolerance are also affected by accli-mation time. Survival above 30 °C(Lewis and Hettler 1968) and below 5'C (Lewis 1965) was progressivelyextended by acclimation temperaturescloser to test values, suggesting thatrapid changes to extreme temperaturesare more likely to be lethal thanprolonged exposure to slowly changingvalues. Winter shutdown of power plantoperations may result in rapidtemperature decreases near the effluentdischarge area. Mortality of juvenileAtlantic menhaden to a temperaturedecrease of 10 °C (from 15 to 5 C) wasless at rates of decrease of 6.7 °C/hor lower than at faster rates. Wintermenhaden kills can be minimized byreducing the rate of decrease as thepower plant discharge is shut down(Burton et al. 1979).

Hettler and Colby (1979) demon-strated that photoperiod at leastpartly explains variation in resis-tance to heat stress. Median lethaltime increased linearly with photo-period at 34 *C. They also speculatedthat it may be important to othertypes of physiological stress. Lewisand Hettler (1968) observed increasedsurvival of juveniles at 35.5 'C withincreased dissolved oxygen (DO)saturation. Burton et al. (1980)reported a mean lethal DO concentra-tion of 0.4 mg/l , but warned againstinterpretation of this value as"safe," in view of the interactivenature of environmental factors.Westman and Nigrelli (1955) observedmass mortalities from gas embolismonly in areas with highly variablesalinity and organic pollutionsufficiently severe to make shellfishunfit for human consumption. Lewisand Hettler (1968) observed decreased

14

survival at high temperatures by fishaffected by gill parasites. Theinteraction of environmental factorsmust be considered when one defineshealthy ranges for-an organism.

Substrate and System Features

The association of the Atlanticmenhaden with estuarine and nearshoresystems during all phases of its lifecycle is well documerted. It is evi-dent that young menhaden require thesefood-rich waters to survive and grow,and the fishery is concentrated nearmajor estuarine systems (June 1961a).Filling of estuarine wetlands, inaddition to exacerbating extremes inenvironmental conditions, has physi-cally limited the nursery habitatavailable to Atlantic menhaden andother estuarine-dependent species.The relative importance, however, ofdifferent habitat types (i.e., sounds,channels, marshes) and salinityregimes has received little detailedattention.

Environmental Contaminants

lack of significant differencesbetween areas within years, althoughthis may have been due to the samplingregime. They speculated that PCBlevels have remained somewhat highbecause of leakage from sourcesestablished prior to regulation andcontinued allowance of limitedspecialty uses. Menhaden oil productscarry the highest concentrations ofsuch non-polar compounds and somesamples contained levels in excess ofUnited States Food and Drug Admini-stration temporary tolerances, as of1977. Warlen et al. (1977) demonstra-ted that 1"4C-DDT uptake by Atlanticmenhaden is dose-dependent, with anassimilation value between 17% and27%. Application of their model . tofield data suggested that uptake wasby way of plankton and detritus.Little information exists about thetoxicity of contaminants to Atlanticmenhaden.

Other Factors

The seasonal depth distributionof Atlantic menhaden is tied to migra-tion patterns. Some fish occur year-round in depths of 1 to 200 m (3 to656 ft). The role of turbidity inAtlantic menhaden biology apparentlyhas not been studied. Blaber andBlaber (1980) proposed that gradientsof turbidity, nutrients, and salinitycould provide cues that enable fry tolocate estuarine nursery areas alongthe Australian coast. The "seeking"of turbid zones is probably related todifferential mortality linked to foodsupply and predation (Blaber andBlaber 1980; Norcross and Shaw 1984).

In a study of chlorinated hydro-carbon residues in menhaden fisheryproducts from the Atlantic and Gulf ofMexico, Stout et al. (1981) showedthat overall levels have decreasedsince the late 1960's, although signi-ficant differences between years forlevels of polychlorinated biphenyls(PCB's) in the South Atlantic Regionand for dieldrin in the MiddleAtlantic Region could not be demon-strated. There was also a general

15

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20

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Life history ofeastern Gulf of

Am. Fish. Soc.

23

$0277: -10

REPORT 00CUM.ENTA7TON 1. R[PS=Rt NO.

PAGZ Biological Report 82(11.108)*

Species Profiles: Life Histories and Environmental Requirements of August 1989Coastal Fishes and Invertebrates (Mid-Atlantic)--Atlantic Menhaden L

f L PJionn~Q•n~ C'gecI, a.•iII en., Nl.I7. A ihas

S. Gordon Rogers and Michael J. Van Den AvyleGeorgia Cooperative Fish and Wildlife Research Unit _

School of Forest ResourcesI C-'•,,,•,c•) ,CG No.

University of GeorgiaAthens, GA 30602

IL Zeo..o..n 0C...,i.fo PA..... s Ad,.q.

National Wetlands Research Center U.S. Army Corps of Engineers IL 7of Ao"'t" L*P.11.4 C-yMa

Fish and Wildlife Service Waterways Experiment StationU.S Department of the Interior P.O. Box 631Washington, DC 20240 Vicksburg, MS 39180

*U.S. Army Corps of Engineers Report No. TR EL-82-4

L Abstract (U.-t: 200 a-oC)

Species profiles are literature summaries of the life history, distribution, and environ-mental requirements of coastal fishes and invertebrates. Profiles are prepared to assistwith environmental impact assessment. The Atlantic menhaden (Brevoortia tyrannus) is animportant commercial fish along the Atlantic coast. In the South Atlantic Region, Atlanticmenhaden spawn during winter in continental shelf waters. Adults then move inshore andnorthward in spring; some move into estuaries as far as the brackish-freshwater boundary.Atlantic menhaden larvae in the South Atlantic Region enter estuaries after 1 to 3 monthsat sea. Young fish move into the shallow regions of estuaries and seem to prefer vegetatedmarsh habitats. Atlantic menhaden are size-selective plankton feeders as larvae, andfilter feeders as juveniles and adults. Due to their large population size, individualgrowth rates, and seasonal movements, Atlantic menhaden annually consume and redistributelarge amounts of energy and materials. They are also important prey for large game fishessuch as bluefish (Pomatomus saltatrix), striped bass (Morone saxatilis), and bluefin tuna(Thunnus thynnus). The Atlantic menhaden is associated with estuarine and nearshoresystems during all phases of its life cycle. Young menhaden require these food-richhabitats to survive and grow. Destruction of estuarine wetlands has decreased nurseryhabitat available to Atlantic menhaden and other estuarine-dependent species.

Estuaries Feeding habitsFishesGrowth

Atlantic menhaden FisheriesBrevoortia tyrannusHabi tatSpawning

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As the Nation's principal conservation agency, the Department ofthe Interior has responsibility for most of our nationally ownedpublic lands and natural resources. This includes fostering thewisest use of our land and water resources, protecting our fishand wildlife, preserving the environmental and cultural values of ournational parks and historical places, and providing for the enjoy-ment of life through outdoor recreation. The Department assessesour energy and mineral resources and works to assure that theirdevelopment is in the best interests of all our people. The Depart-ment also has a major responsibility for American Indian reservationcommunities and for people who live in island territories under U.S.administration.

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U.S. DEPARTMENT OF THE INTERIORFISH AND WILDLIFE SERVICE

TAKE PRIDEin America

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UNITED STATESDEPARTMENT OF THE INTERIOR

FISH AND WILDLIFE SERVICENational Wetlands Research Center

NASA-Slidell Computer Complex1010 Gause Boulevard

Slidell, LA 70458

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