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Fws/Usace - Species Profile: Atlantic Menhaden
ML072060570
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Site: Oyster Creek
Issue date: 08/01/1989
From: Moran D, Shannon Rogers, Vandenavyle M
Univ of Georgia, US Dept of Interior, Fish & Wildlife Service, US Dept of the Army, Corps of Engineers
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Davis J NRR/DLR/REBB, 415-3835
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TR EL-82-4 82(11.108)
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Coastal Ecology Group Fish and Wildlife Service Waterways Experiment Station U.S. Department of the Interior U.S. Army Corps of Engineers jA-y Fw$MO\(Q$~LC e..

Biological Report 82(11.108)

TR EL-82-4 August 1989 Species Profiles: Life Histories and Environmental Requirements of Coastal Fishes and Invertebrates (Mid-Atlantic)

ATLANTIC MENHADEN by S. Gordon Rogers and Michael J. Van Den Avyle Georgia Cooperative Fishery Research Unit School of Forest Resources University of Georgia Athens, GA 30602 Project Officer David Moran National Wetlands Research Center U.S. Fish and Wildlife Service 1010 Gause Boulevard Slidell, LA 70458 Performed for Coastal Ecology Group Waterways Experiment Station U.S. Army Corps of Engineers Vicksburg, MS 39180 and U.S. Department of the Interior Fish and Wildlife Service Research and Development National Wetlands Research Center Washington, DC 20240

This series may be referenced as follows:

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

This profile may be cited as follows:

Rogers, S. G., and M. J. Van Den Avyle. 1989. Species profiles: life histories and 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 profiles are designed to provide coastal managers, engineers, and biologists with a brief comprehensive sketch of the biological characteristics and environmental requirements of the species and to describe how populations of the species may be expected to react to environmental changes caused by coastal development. Each profile has sections on taxonomy, life history, ecological role, environmental requirements, and economic importance, if applicable. A three-ring binder is used for this series so that new profiles can be added as they are prepared.

This project is jointly planned and financed by the U.S. Army Corps of Engineers and the U.S. Fish and Wildlife Service.

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

Information Transfer Specialist National Wetlands Research Center U.S. Fish and Wildlife Service NASA-Slidell Computer Complex 1010 Gause Boulevard Slidell, LA 70458 or U.S. Army Engineer Waterways Experiment Station Attention: WESER-C Post Office Box 631 Vicksburg, MS 39180 iii

CONVERSION TABLE Metric to U.S. Customary Multiply To Obtain mill-imeters (mm) 0.03937 inches centimeters (cm) 0.3937 inches meters (iM) 3.281 feet meters (m) 0.5468 fathoms kilometers (km) 0.6214 statute miles kilometers (km) 0.5396 nautical miles 2

square meters (m )

2 10.76 square feet square kilometers (km ) 0.3861 square miles hectares (ha) 2.471 acres liters (1) 0.2642 gallons cubic meters (m3 ) 35.31 cubic feet 3

cubic meters (m ) 0.0008110 acre-feet milligrams (mg) 0.00003527 ounces grams (g) 0.03527 ounces kilograms (kg) 2.205 pounds metric tons (t) 2205.0 pounds metric tons (t) 1.102 short tons 3.968 kilocalories (kcal) British thermal units Celsius degrees ('C) 1.8( 0 C) + 32 Fahrenheit degrees U.S. Customary to Metric inches 25.40 millimeters inches 2.54 centimeters feet (ft) 0.3048 meters fathoms 1.829 meters statute miles (mi) 1.609 kilometers nautical miles (nmi) 1.852 kilometers square feet (ft 2 ) 0.0929 square meters square miles (mi 2 ) 2.590 square kilometers acres 0.4047 hectares gallons (gal) 3.7a5 liters 3

cubic feet (ft ) 0.02831 cubic meters acre-feet 1233.0 cubic meters ounces (oz) 28350.0 milligrams ounces (oz) 28.35 grams pounds (lb) 0.4536 kilograms pounds (lb) 0.00045 metric tons short tons (ton) 0.9072 metric tons British thermal units (Btu) 0.2520 kilocalories Fahrenheit degrees (*F) 0. 5556 (-F 32) Celsius degrees iv

CONTENTS Page PREFACE .................................................... iii CONVERSION TABLE .................. ......................... IV ACKNOWLEDGMENTS .......................................................... vi NOMENCLATURE/TAXONOMY/RANGE ............................................. 1 MORPHOLOGY/IDENTIFICATION AIDS ...................................... 1 REASON FOR INCLUSION IN SERIES ........................................... 3 LIFE HISTORY... ......... ....................... 3 Adult Migration and Spawning ........................................... 3 Fecundity .............................................................. 4 Eggs and Larvae ......................................................... 4 Juveniles and Adults................................................... 6 GROWTH CHARACTERISTICS ................................................... 7 THE FISHERY ....... ...................................................... 9 History ................................................................ 9 The Catch ......................................... ..................... 9 Management ............................................ ................. 10 Subpopulations ......................................................... 11 ECOLOGICAL ROLE .......................................................... 12 ENVIRONMENTAL REQUIREMENTS ............................................... 13 Temperature, Salinity, and Dissolved Oxygen ............................ 13 Substrate and System Features .......................................... 14 Environmental Contaminants ............................................. 15 Other 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 Smith of the National Marine Fisheries Service, Beaufort, North Carolina, and by Walter Nelson, National Marine Fisheries Service, Miami, Florida.

vi

Figure 1. Atlantic menhaden.

ATLANTIC MENHADEN NOMENCLATURE/TAXONOMY/RANGE MORPHOLOGY/IDENTIFICATION AIDS Scientific name: Brevoortia tyrannus Branched dorsal rays, 13-18; (Latrobe) branched anal rays, 16-22; pectoral Preferred common name ......... Atlantic rays, 14-18; pelvic rays, 7; gill fil-menhaden (Robins et al. 1980; aments, 51-66; lateral line scales, Figure 1) 40-50; ventral scutes, 29-34; verte-Other common names: pogy, moss- brae, 44-49. Body oblong and com-bunker, bunker, fat-back, shad, pressed with a thin belly wall; scales bug-mouth large, coarse, with long slender Class ..................... Osteichthyes pectinations, strongly overlapping and Order ..................... Clupeiformes in regular rows; predorsa! scales on Family ............ Clupeidae (herrings) either side of median line enlarged; prominent radiating opercular stria-Geographic range: Temperate coastal tions; and pelvic fin rounded with waters from Nova Scotia southward, innermost and outermost rays about to Jupiter Inlet, Florida (Dahlberg equal in length (Hildebrand 1963; 1970). Atlantic menhaden are sea- Dahlberg 1970, 1975).

sonally abundant in the Mid-Atlantic

  • Region (Figure 2). Concentrations Color in life: green, brown, or of age 0 fish occur in inshore blue-gray, darker on dorsal surface.

estuarine waters along the entire A dark humeral spot may be followed Atlantic seaboard. posteriorly by a series of smaller 1

NEW YORK 0

ATLANTIC OCEAN 0

MILES 0 50 100 0 so 100 KILOMETERS 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 upon in estuarine zones and nearshore shelf capture. Brevoortia tyrannus can be waters northward from Chesapeake Bay.

distinguished from B. smithi (yellow- Due to the species' great abundance, fin menhaden, the only other North extensive migration patterns, and American coast species) because B. importance as a prey species, the tyrannus has a frontal groove, larger Atlantic menhaden influences the and coarser scales in regular rows conversion and exchange of energy and (therefore, lower scale-related organic matter within biological counts), pointed (versus rounded) systems throughout its range (Peters scale pectinations, a row of lateral and Schaaf 1981; Lewis and Peters spots behind the humeral spot, more 1984).

gill filaments on the ceratobranchial arch, rounded pelvic fins, and opercu-lar striations. The caudal fin of B. LIFE HISTORY smithi is bright yellow, whereas the caudal fin of B. tyrannus is not.

Fresh B. tyrannus may have a darker Terminology used to describe life anal fin and more body mucus than does history stages conforms to that used B. smithi. Atlantic menhaden can be by Lewis et al. (1972) and Moyle and distinguished from F1 hybrid Atlantic Cech (1982).

menhaden (Brevoortia smithi x B.

tyrannus) by its longer frontal groove, lateral spots (absent to few Adult Miqration and Spawning in hybrid), and a rounded ventral fin.

Dahlberg (1970, 1975) provided addi- Knowledge of timing and location tional measurements and descriptions of spawning has been mainly obtained of qualitative characters. Jones et from collections of adult females that al. (1978) gave detailed descriptions were spent or contained maturing ova of Atlantic menhaden developmental (Higham-and Nicholson 1964; June 1965; stages (egg through adult). Hettler Dahlberg 1970) as well as from collec-(1984) gave meristic and morphometric tions of eggs and larvae (Reintjes descriptions of Atlantic menhaden 1961, 1968; Herman 1963; Kendall and larvae and juveniles. Reintjes 1975; Ferraro 1980a,b; Judy and Lewis 1983). Data on movement and population age or size structure have REASON FOR INCLUSION IN SERIES been obtained from distribution of purse-seining effort (Roithmayr 1963);

frequencies of lengths, weights, and Atlantic menhaden constitute ages in catches (June and Reintjes about 25% to 40% of the combined 1959;. June 1972; Nicholson 1971, annual landings of Atlantic coast and 1972); and returns from extensive Gulf of Mexico menhaden species, which tagging experiments (Dryfoos et al.

collectively comprised the largest 1973; Kroger and Guthrie 1973; commercial fishery by weight and Nicholson 1978). Atlantic menhaden eighth largest in dollar value in undergo extensive north-south seasonal 1984 in the United States (NMFS migrations and inshore-offshore 1985). They are important prey for movements along the Atlantic seaboard.

many other fish species and are Schools are composed of fish of seasonally important components of similar size and age. Migration estuarine and shelf fish assemblages. patterns are also related to spawning Atlantic menhaden depend on habitats habits, and some spawning occurs every along the entire eastern seaboard and month of the year.

adjacent shelf waters throughout their life cycle and use estuaries as Atlantic menhaden of all ages nursery areas. Some spawning occurs congregate off North Carolina from 3

November to March and some spawn there Table 1. Estimated fecundity of in shelf waters that are 100 to 200 m Atlantic menhaden at different ages deep, probably within 70 m of the (from Dietrich 1979).

surface (Reintjes and Pacheco 1966).

All eggs of Atlantic menhaden collected by Judy and Lewis (1983) Number of eggs per female were taken at depths of 10 m or less. (thousands)

The spawning is heaviest off Cape Age Mean Range N Lookout, North Carolina, from December through February. Adults then move inshore and northward in spring and 1 115.8. 26.5 - 250.7 21 stratify by age and size along the Atlantic seaboard. Adult menhaden II 177.4 39.2 - 368.8 34 have been collected from estuaries, and some move as far inland as the 1I1 302.8 127.7 - 458.3 33 brackish-freshwater boundary. The oldest and largest fish migrate the IV 308.6 142.7 - 514.0 12 farthest, some moving as far north as the Gulf of Maine.. Adults that remain V 568.4 --- 1 in the South Atlantic Region move southward and reach northern Florida by fall. Representatives of all age classes return to the shelf waters of Nicholson (1964) gave the following the South Atlantic Region in late equation for the estimation of fecun-fall. dity (F = ova per fish; FL = fork length, mm):

During migration northward in spring, spawning occurs progressively In F = 7.2227 + 0.0176 FL closer inshore; by late spring, some fish spawn within coastal embayments r2 = 0.726.

of the North Atlantic Region. There are definite spring and fall spawning Dietrich (1979) gave the following peaks in the Middle and North Atlantic equations (W = wet body weight less Regions, and some spawning occurs weight of ovaries, g; A = age, years; during the winter in the shelf waters FL = fork length, mm):

of the Mid-Atlantic Region. Temporal and spatial segregation of spawning F = 488 W activity provides a mechanism for the r2 = 0.916 existence of races (= subpopulations).

Higham and Nicholson (1964) and Schaaf F = 92,592 A (1979) have speculated that a female r 2 = 0.879 may spawn more than once in a season.

In F = 8.6463 + 0.0120 FL r2 = 0.871.

Fecundity Eggs and Larvae Higham and Nicholson (1964) reported values of 38,000 to 631,000 Eggs of the Atlantic menhaden are ova per fish, and June (1961a) gave pelagic and have been reported to values of 40,000 to 700,000 ova per hatch in 2 days at unspecified temper-fish; estimates depend on the size of atures (Kuntz and Radcliffe 1917), 2.9 the fish. Dietrich (1979) reported days at 15.5 'C (Ferraro 1980a), and fecundities of 116,000 to 568,000 at 2.5 to 2.9 days at an average ova per fish at age I to age V, temperature of 15.5 CC (Hettler respectively (Table 1). Higham and 1981).

4

Survival of laboratory reared al. 1980). Massmann et al. (1954)

Atlantic menhaden embryos to hatching reported that the abundance of pre-is very low (2% to 45%); 49% to 94% of juveniles was higher above than below mortality occurs before blastopore the brackish-freshwater boundary, and closure (Ferraro 1980a). Rogers et al. (1984) showed that this pattern persists during high river Atlantic menhaden larvae begin discharge. Massmann et al. (1954),

feeding on individual zooplankters Rozas and Hackney (1984), and Rogers (Reintjes and Pacheco 1966) about 4 et al. (1984) provided evidence that days after hatching when the yolk sac prejuvenile Atlantic menhaden select is almost absorbed, the eyes are tidewater areas that are fresh or of pigmented, and the mouth is func- low salinity. Only fish of prejuve-tional (Hettler 1981). Larvae in the nile lengths were resident in low South Atlantic Region enter estuaries salinity river shoals and in inter-after 1 to 3 months at sea (Reinties tidal creeks (Lewis et al. 1972).

1961) at fork lengths (FL) of 14 to This phenomenon persisted for about 4 34 mm (Reintjes and Pacheco 1966); months (Rogers et al. 1984).

fish longer than 30 mm FL are then already metamorphosing to the pre- A "critical period" of survival juvenile morphology (Lewis et al. in young fishes was first defined by 1972). This migration into estuaries Hjort (1914) and discussed for clupei-occurs from May through October in form fishes by Schumann (1965) and the North Atlantic Region, October May (1974). Menhaden, like most through June in the Mid-Atlantic, and fishes with high fecundity and little December through May in the South parental care, hatch in an undeveloped Atlantic Region (Reintjes and Pacheco state. Such fish typically rely on 1966). As they grow, the larvae energy contained in the yolk-sac for probably feed on progressively larger 4-5 days after hatching, after which zooplankters (Kjelson et al. 1975). they are sufficiently developed to more efficiently feed on an external Young fish move into the shallow food supply (Schumann 1965). Feeding portions of estuaries including river of the youngest Clupea, Engraulis, shoals and the heads of small tidal and Sardinops larvae depends largely creeks (Massmann 1954; Massmann et al. on food availability; fish will eat 1954; June and Chamberlin 1959; to capacity in the presence of high Pacheco and Grant 1965; Wilkens and food concentrations and starve in Lewis 1971; Lewis et al. 1972; the absence. of high concentrations Weinstein 1979; Weinstein et al. because they are unable to move about 1980; Rozas and Hackney 1984; Rogers to search for food. A routine search et al. 1984). They apparently prefer pattern is initiated only after an Spartina, Juncus, and the vegetation encounter with or capture of a food typical of fresh tidal marshes and particle. Given the heterogeneity in swamps (Taxodium, Typha, Nyssa) distribution of pelagic plankters and Peltandra, Rumex, Sagittaria, Zizania) the inability of many clupeiform over vegetated habitats in open water fishes to cope with low food concen-(Adams 1976; Weinstein and Brooks trations, menhaden probably have a 1983). While in estuaries, Atlantic critical period of larval survival.

menhaden grow and metamorphose through Year-class strength may be partly a prejuvenile stage into juveniles. determined during this period. This problem is most likely to occur when Several studies have reported larvae are spawned offshore or swept abundances of young menhaden that were offshore after having been spawned higher in portions of estuaries with nearshore. Individual larval condi-the lowest salinity <5 ppt (Lewis et tion factors (weight-length ratios) al. 1972; Weinstein 1979; Weinstein et increase rapidly when the fish enter 5

K an estuary (Lewis and Mann 1971). Beaufort, North Carol i.na ; pers.

Metamorphosis is not totally dependent comm.).

on low salinity; Hettler (1981) reared Atlantic menhaden from eggs to Survival of Atlantic menhaden to juveniles using water with a high age I has been estimated by comparing salinity. However, metamorphosis estimates of population size of age I rarely is successful outside the fish (based on a virtual population food-rich, low-salinity estuarine analysis that incorporated data from zones (Kroger et al. 1974). commercial landings) with number of eggs predicted to have been spawned in previous years. Estimates of No data are available that link recruits per million eggs spawned survival at yolk sac absorption to have ranged from 27 to 159 (Nelson et year-class strength or that enable al . 1977) and from 78 to 282 quantitative estimates of mortality (Dietrich 1979).

from predation or starvation (May 1974). Minimum food concentration for inception of feeding activity is not Juveniles and Adults known, and survival curves do not exist for larval Atlantic menhaden. Metamorphosis marks a change in Nelson et al. (1977), however, feeding mode from capturing individual developed an environment-recruit model zooplankton to filter-feeding (June in an attempt to explain variation in and Carlson 1971; Durbin and Durbin year-class strength. Since larval 1975). This shift is accompanied by a menhaden are thought to depend largely loss of teeth,, an increase in the on wind-driven (Ekman) transport to number and complexity of gill rakers, reach estuarine nursery grounds and an increase in the complexity and (particularly in the Middle and South musculature of the digestive tract Atlantic Regions), the modelincorpo- (June and Carlson 1971). Prejuveniles rated four variables: (1) the known are somewhat intermediate in feeding spawning times and locations; (2) wind mode (June and Carlson 1971) and body vectors specific for year, time, and structure (June and Carlson 1971; location; (3) year- and time-specific Lewis et al. 1972).

discharges of major tributary systems; and (4) the minimum sea surface Juveniles begin congregating in temperature at the mouth of Delaware dense schools as they leave shoal Bay. A survival index was calculated areas. Most emigrate from estuaries as the ratio of observed recruitment from August through November (earliest to the fishery (age I) to that predic- in the North Atlantic Region) at ted by a Ricker (1954) spawner-recruit lengths of 55 to 150 mm FL (June (density-dependent) model. The magni- 1961a) or 55 to 140 mm total length tude of the index "should reflect (TL) (Nicholson 1978). Nicholson those environmental (density indepen- stated that most emigrants are 75 to dent) effects which influence survival 110 mm TL. As judged by the results of menhaden from the time of spawning of extensive tagging, many age 0 fish to the time of recruitment to the migrate southward along the North fishery at age I" (Nelson et al. Carolina coast in late fall and 1977). The model explained 84% of the early winter (Nicholson 1978). Fish variation in the survival index for in the southernmost portion of the the years investigated (1961-71); South Atlantic Region, however, Ekman transport was the principal showed less offshore migration component. The correlation has not (Dahlberg 1970), and tagging results been statistically significant in indicated that juveniles leaving the recent years (D. Vaughan, National estuaries of the South Atlantic Region Marine Fisheries Service (NMFS), and the North Atlantic Region may not 6

move far north or south during their GROWTH CHARACTERISTICS first year (Nicholson 1978). Larvae entering estuaries late in the season may remain in the estuary one addi- Growth rates vary among years and tional year and emigrate at. age I. localities throughout the species' Some juveniles and adults are found in range (June and Reintjes 1959, 1960; sounds and bays along the South June 1961b; June and Nicholson 1964; Atlantic Region during mild winters Nicholson and Higham 1964a, 1964b, (June 1961a). Fish leaving estuaries 1965; Nicholson 1975; Reish et al.

along the entire Atlantic seaboard 1985). The age of Atlantic menhaden eventually disperse throughout most of can be determined from annual scale the geographic range (Nicholson 1978). markings (McHugh et al. 1.959; June and Roithmayr 1960; Kroger et al. 1974).

Most Atlantic menhaden reach Fish of the same age are progres-maturity by the end of their second sively larger in more northerly fish-full year. About 10% were found to eries (Nicholson 1978; Reish et al.

be capable of spawning at age I (late 1985), but mature at smaller sizes in in the year), and 90% at age II more southerly areas. Minimum size at (Higham and Nicholson 1964). Fish of maturity was 180 mm FL in the South all ages, however, are found in the Atlantic and 210 mm FL in the Middle migrating schools. Although Atlantic Atlantic (Higham and Nicholson menhaden can live 8 to 10 years (June 1964). There is evidence that growth and Roithmayr 1960), fish older than rates have changed in response to age IV had been rare in the commercial fishing pressure: fish of the same age catch. As stocks rebuild, however, were larger in the late 1960's and age V and VI fish are becoming more early 1970's than in the late 1950's common and may be locally abundant in and early 1960's (Nicholson 1975).

the North Atlantic and North Carolina Reish et al. (1985) indicated that fall fishery (Powers 1984). growth rates do not depend upon fish abundance. Atlantic menhaden in years of high abundance probably are smaller Adult feeding behavior is than Atlantic menhaden in years of low affected by food availability (Durbin abundance because the former were and Durbin 1975; Durbin et al. 1981). smaller at the time of recruitment, Swimming speed is increased at higher not because of any difference in food concentrations, and associated growth rates after recruitment.

energetic costs rise exponentially.

Modeling studies have suggested that Growth has been shown to be allo-Atlantic menhaden maximize growth metric in larval, prejuvenile (Lewis rate, not efficiency (Durbin and et al. 1972), juvenile (Lewis et al.

Durbin 1983), and that efficiency of 1972; Epperly 1981), and adult stages.

dietary assimilation changes seasonal- Reish et al. (1985), however, reported ly with the quality of available food. that Atlantic menhaden exhibited The Atlantic menhaden has behavioral, growth that was close to isometric, physiological, and morphological although growth appeared to become adaptations for an active migratory allometric with increasing age. Three existence in waters with pronounced different "stanzas" of growth in young seasonal and spatial variation in food menhaden were reported by Lewis et al.

abundance. It has large lipid (1972), with inflection points reserves that are seasonally assimi- at 30 and 38 mm TL (70 and 469 mg lated (Durbin and Durbin 1983), a body weight). These points served as the shape adapted for continuous swimming, basis for their division of the life and copious body mucus (Dahlberg history stages. Balon (1984) pointed 1970). out that these are functional, not 7

arbitrary, divisions. Lewis et al. spawned and entered the estuary.

(1972) cited an unpublished manuscript Young of the next year class arrived that stated that the relationship in the spring only 20 to 30 mm TL between length andweight is similar shorter than the smallest fish of the for juveniles and adults. Length- previous year class. These factors, weight conversions can be made using combined with differences in larval the appropriate equation in Table 2. growth rates (Lewis et al. 1972; Kro-ger et al. 1974) and latitudinal dif-Atlantic menhaden growth begins ferences in growing season, probably in spring and ends in fall, as the explain the observed differences in water temperature crosses an approxi- sizes of fish of the same "age" within mate 15 *C threshold (Kroger et al. a single year's catch. See Durbin 1974). At Beaufort Inlet, North Caro- and Durbin (1983) and the section on lina, age 0 fish ranged from 40 to 185 Life History for discussions of mm TL at the end of the growing sea- growth in relation to feeding, son, depending on when they were environment, and body morphology.

Table 2. Weight-length regressions for Atlantic menhaden; loge weight = a + b (log e length).

Types of measurement 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 3.145 and offspring 2 Middle Atlantic2 spawners and g SL -11.037 3.103 offspring South Atlantic 2 spawners and g SL -10.579 2.995 offspring All spawners, 3 for the fishery g SL -12.075 3.215 as a whole

'Lewis 2 et al. 1972.

Epperly 1981.

3 Douglas Vaughan, National Marine Fisheries Service, pers. comm.

8

Atlantic menhaden reach lengths recently included primarily age I and of about 500 mm TL and weights of over II fish. During the summer fishery in 1,500 g at ages of 8 to 10 years. the South Atlantic Region, fish caught Cooper (1965) collected an 8-year-old have been mostly ages I and 11 except that measured 470 mm TL and weighed in 1984 when a large number of age 0 1,674 g. fish were caught; the north Florida landings have been composed mostly of age I fish. Concurrently,' fish in the THE FISHERY, Chesapeake Bay area and the southern portion of the Mid-Atlantic Region are History also age I and II, although they are longer and heavier on the average.

The Atlantic menhaden fishery was Some fish of ages III and IV are first established in the late 1600's present in an early spring pound-net and early 1700's to obtain fish for fishery in Chesapeake Bay. Most of agricultural fertilizer (Frye 1978). the fish caught in the northern In the early 1800's, an industry was portion of the Mid-Atlantic Region are developed to obtain oil f'rom of ages 11 and III, the age II fish menhaden (Goode 1879; Goode and Clark being larger than those to the south.

1887), and by 1869 there were 90 For the entire Atlantic menhaden reduction plants in North Carolina fishery, the average percentage of alone (June 1961a). Today this numbers of age I and 11 fish between species contributes 25% to 40% of the 1955 and 1984 was 78.4%; the range was landings in the largest commercial 51.4% (in 1961) to 95.9% (in 1970).

fishery (by weight, Brevoortia Numbers of age 0 fish composed 25% or species) in the United States. Annual more of the catch for the entire landings for 1979 to 1981 averaged fishery in 1955, 1966, 1979, 1981, about 400,000 metric tons and $38 1983, and 1984 (D. Vaughan, National million in market value (NMFS 1980, Marine Fisheries Service; pers.

1981, 1982, 1983, 1984, 1985). Plants comm.). -The north Atlantic fishery that process Atlantic. menhaden operates from mid-June through October products currently operate from Maine and primarily exploits fish of age III to Florida. About 96% *to 918% of the or older. The purse seine fishery catch is sold to livestock and cos- north of Cape Hatteras is over by late metic interests as fishmeal, soluble November. Age 0 fish begin to be proteins, and oils; the rest is used vulnerable to the fishery during late in pet food products and as fish bait fall and winter from Chesapeake Bay (NMFS 1980, 1981, 1982, 1983, 1984, south. The North Carolina fall 1985). Most of the landings are made fishery is composed of fish of all age with purse seines. Federal efforts to classes; age 0 fish have predominated collect data for management of Atlan- since 1971, except in 1980 and 1982.

tic menhaden began in 1955 (June 1957). Atlantic menhaden stocks were drastically reduced during the 1960's.

Annual landings dropped from 671,400 The Catch metric tons in 1955 to about 200,000 metric tons per year from 1966 through The Atlantic menhaden fishery has 1969 (Nelson et al. 1977). As the two annual phases: a summer and fall population size decreased, the age fishery from Maine to northern structure also changed. Fish older Florida, and an intensive fall and than age III became scarce and fish winter fishery off North Carolina older than age IV were practically between Cape Lookout and New River non-existent even in the North Inlet (June 1961a; Nicholson 1978). Carolina fall-winter fishery. Many Landings for the entire fishery have northern processing plants closed 9

down -- especially those in the New State agencies; only the states have England area, where the fishery final regula.tory power.

depended on fish of ages III and IV (Henry 1971; Nicholson 1975). Age I The stocks have generally been and II fish constituted the bulk of unmanaged. Nelson et al. (1977) the landings and age 0 became more suggested that the fishery is somewhat important (Nicholson 1975). The self-requlating in that reduced stocks began to recover in the early catches bring about reduced effort and 1970's, when age III fish again plant closures, allowing the stocks to appeared in North Atlantic catches. recover. They stress that proper The first significant Maine landings management practices could reduce the (3,100 metric tons) since the 1960's chance of repeating the mistakes of occurred in 1973 (NMFS 1973, 1974, the past and prevent a crash in the 1975; Nelson et al. 1977). North fishery. In addition, Schaaf (1975)

Atlantic (from Cape Cod, Massachu- pointed out that "allowing the fishery setts, to Cape Breton, Nova Scotia) to be controlled . . . by the econo-landings in 1929-71 correlated mics of free market competition strongly with 3-year-lagged local assures that (1) the average profit water temperatures and mixing factors for the whole industry will be zero

%for St. Lawrence River inputs * . . and (2) there is no mechanism to

'(Sutcliffe et al. 1977). These North provide for protection of the Atlantic fish, though vulnerable resource, since if either costs go throughout the fishery, may be a down or value goes up new effort can unique biological stock. Catches afford to enter the fishery and continued to improve into the early eventually may exceed the BBEP 1980's; however, the size of the [Biological Break-Even Point, a point reproductive stocks (ages III and IV) at which the fishery collapses]."

remained low (Atlantic Menhaden Schaaf (1975) also warned that the Management Board [AMMB] 1981; NMFS level of effort when the fishery 1983, 1984, 1985). Heavy exploitation collapsed in the 1960's might be results in smaller and fewer fish maintained during the mid-1970's even (June 1972) and higher catchability if catches dropped to 200,000 metric coefficients (Schaaf 1979). The tons per year because product prices implications of these phenomena are were higher at that time. He urged not fully understood; however, a recent implementation of a flexible quota model showed that pollution stress may system coordinated by the Atlantic greatly reduce first-year survival rate States Marine Fisheries Commission (Schaaf et al. 1987). (ASMFC) that would allow the stocks to continue to rebuild while effort Management regulation mechanisms were studied.

More than 50% of the annual The management option endorsed by landings of Atlantic menhaden are. from the ASMFC, Option 7 of the Atlantic within State territorial waters, Menhaden Advisory Committee, proposed mostly from the Chesapeake Bay area the following closing dates to protect (R. B Chapoton, NMFS, Beaufort, North age 0 recruits: the week ending Carolina; pers. comm.). This fact, between October 4 and 10 for the North combined with the migratory nature of Atlantic, the week ending between the species and the dependence of October 11 and 17 for the Mid-northern fisheries on. escapement of Atlantic, the week ending between age I and II fish from fisheries in November 8 and 14 for Chesapeake Bay, the South Atlantic Region and and the week ending between December Chesapeake Bay (Nicholson 1978), makes 13 and 19 for the South Atlantic.

regulation a _compromise, situation This closure would primarily affect between the industry and Federal and the North Carolina late-fall fishery.

10

To be effective, the regulation still (1977) likewise averaged 419,000 needs to be implemented in North metric tons per year (1961-71) and is Carolina; it has been implemented proposed to offer a way of "fine-already in six states (Vifrginia and tuning" predicted catch on a yearly north). National Marine Fisheries basis by constantly updating yield Service personnel 'have analyzed the estimates. The vulnerability of the effects that Option 7 would have by Atlantic menhaden fishery. to fluctua-using data for 1976-82. Increases in tions in year class strength was first yield-per-recruit ranged from 0.4% to pointed out by June (1961a). It has

>10% and varied with strength of the since been stressed that the mainte-targeted age 0 year class and the nance of a healthy stock of spawning-timing of arrival at the North age fish should be a primary concern Carolina fall fishery. These esti- of management (Schaaf and Huntsman mates may be low because of sampling 1972; Schaaf 1975; Nelson et al. 1977; errors, but would be substantially Vaughan 1977). Good stocks of greater only in years with very spawning-age fish would bring multiple abundant age 0 cohorts that arrive at benefits, including higher reproduc-the North Carolina fall fishery tive potential (decreasing the effects after the mid-December closing date of years with poor recruitment),

(D. Vaughan, pers. comm.). This decreased vulnerability to weak year management strategy is an extension of classes, and increased weight of NMFS recommendations made since the landings due to a higher contribution early years of study. The continued of older fish to the catch. Recent existence of the Atlantic menhaden calculations of MSY remain near fishery despite heavy exploitation is 450,000 metric tons (D. Vaughan, quite unlike that of other well- pers. comm.). I studied clupeid stocks (Schaaf 1979),

and probably is a product of differ- Annual instantaneous natural mor-ences in density-independent phenomena tality was estimated at 0.36 (1955-64)

(e.g., reproductive strategy). In by Schaaf and Huntsman (1972) and at spite of the difficulties involved in 0.42 for the 1955 year class by Nelson managing such a complex resource, the et al. (1977). Their respective esti-prospects of developing a workable mates of annual instantaneous fishing management plan are good, primarily mortality were 0.82 to 2.14 (1955-64) due to the capability of the stocks to and 0.36 for the 1955 year class. A respond to conservation of age O-fish combination of these data yieldstotal (Schaaf 1975). However, the coopera- instantaneous mortality estimates tion of affected states is necessary ranging from 1.18 to 2.56, which to effectively implement such a plan. correspond to total annual mortality rates of 69% to 92% (ages I - VI).

Annual maximum sustained yield Reish et al. (1985) estimated annual (MSY) from historic catch-effort data natural mortality at 0.54 for late has been estimated by Schaaf and juveniles (9-23 months), 0.15 for age Huntsman (1972) at 600,000 metric I (12-23 months), 0.49 for age II tons and by Schaaf (1975) at 560,000 (24-35 months), and 0.52 for age 3+

metric tons. Estimates incorporating (36+ months). Except for the age I the use of a Ricker (1954) spawner- fish, these estimates were comparable recruit (density-dependent survivor- with the annual mortality of 0.52 ship) model were given as 380,000 obtained from mark-recapture data by metric tons per year by Schaaf and Dryfoos et al. (1973).

Huntsman (1972) and averaged 419,000 metric tons per year on the basis of Subpopulations known survivorship in 1961-71 (Nelson et al. 1977). The recruit-environment Because a genetically distinct model developed by Nelson et al. stock can have its own homogenous I1

vital parameters of recruitment, before they enter the estuary do not growth, and mortality (Cushing 1968), exist. After entering the estuary, identification of the stock (=subpopu- Atlantic menhaden larvae appear to be lations) and stock-specific biological extremely selective for prey of cer-traits is necessary for proper manage- tain sizes and species. Larvae from ment. Various authors have proposed the Newport River Estuary, North the existence of two to five Atlantic Carolina, 26 to 31 mm TL (x = 29 mm menhaden. subpopulations on the basis TL), consumed copepods and copepodites of meristic and morphometric compari- of only four taxa, which composed 99%

sons (June 1958, 1965; Sutherland by numbers and volume of their gut 1963; Higham and Nicholson 1964; June contents (Kjelson et al. 1975). These and Nicholson 1964; Dahlberg 1970; prey items, ranging from 300 to 1200.

Epperly 1981). Dahlberg (1970) um in length (ý = 750 um), were eaten reported a distinct subpopulation of despite an abundance of copepod Atlantic menhaden south of Cape nauplii, barnacle larvae, and small Canaveral' in the vicinity of the adult copepods in plankton tows.

Indian River, Florida. Nicholson Larvae that were offered copepods in (1978) stated that the extensive the laboratory ignored all other food north-south migrations, latitudinal items, including Artemia and Balanus stratification by age and size in the nauplii (June and Carlson 1971).

summer, and intermingling of all age Larval menhaden in the Newport River classes south of Cape Hatteras in Estuary, North Carolina, fed primarily winter preclude the existence of more during daylight (Kjelson et al. 1975).

than one stock. Epperly (1981),

however, provided electrophoretic as well as meristic and morphometric data Juvenile and adult Atlantic that indicated significant differences menhaden strain particulates from the between fish spawned in the waters water column with a complex set of north of Long Island, New York, during gill rakers. The rakers can sieve the spring and those spawned in the particles down to 7-9 pm in size fall and winter in the South and (Friedland et al. 1984), including Mid-Atlantic Regions. Other groups -- zooplankton, larger phytoplankton, and fall-spawning fish of the Gulf of chain-forming diatoms. Biochemical Maine and spring-spawning fish of the analyses indicated that the gut Mid-Atlantic Region -- may also be contents of juveniles vary with prey distinct subpopulations, but this availability; reliance on zooplankton aspect has not been fully investi- decreases as the fish move from open gated. Menhaden species hybridize waters to. marshes (Jeffries 1975).

readily (Turner 1969; Dahlberg 1970), Atlantic menhaden may also be capable but Atlantic x yellowfin hybrids have of eating epibenthic materials (Edgar been recorded only as far north as and Hoff 1976). Peters and Schaaf Beaufort, North Carolina (Dahlberg (1981) speculated that the annual 1970). Apparently, hybrids do not phytoplankton and phytoplankton-based occur in the Mid-Atlantic Region. production in east coast estuaries is not sufficient to support the juvenile Atlantic menhaden population during ECOLOGICAL ROLE its residency and that the abundant organic detritus may be eaten. Lewis and Peters (1984) reported that Atlantic menhaden occupy two juvenile Atlantic menhaden in North distinct types of feeding niches Carolina salt marshes ate primarily during their lifetime. They are size- detritus.

selective plankton feeders as larvae and filter feeders as juveniles and The roles of Atlantic menhaden in adults. Data on the food of larvae systems function and community 12

dynamics have received little ENVIRONMENTAL REQUIREMENTS attention. Larvae and juveniles are seasonally important components of Temperature, Salinity, and Dissolved estuarine fish assemblages (Tagatz and Oxygen Dudley 1961; Cain and Dean 1976; Bozeman and Dean 1980). Estimates of Atlantic menhaden occur through a the mean daily ration for larvae range wide range of physicochemical condi-from 4.9% (Kjelson et al. 1975) to 20% tions. Several studies have raised (Peters and Schaaf 1981) of wet body questions about limits of tolerance weight. Assimilation of ingested and optimum conditions. June and energy exceeded 80% for plant and Chamberlin (1959) and Reintjes and animal material (Durbin and Durbin Pacheco (1966) reported that larval 1981). Because of their tremendous menhaden did not enter estuarine numbers, individual growth rates, and waters at temperatures below 3 *C.

seasonal movements, these fish Many studies have noted an affinity of annually consume and redistribute young menhaden for low salinity waters large amounts of energy and materials, (see the Life History section).

including exchanges between estuarine Wilkens and Lewis (1971) speculated and shelf waters. that larval menhaden require low salinity water to metamorphose properly, and Lewis (1966) found that Kjelson et al. (1975) noted that although larvae metamorphosed in.

the copepod taxa preferred by larval salinities of 15 to 40 ppt, one-third menhaden and other species decreased of the juveniles developed slightly from a mean value (2 years) of 81% to crooked vertebral columns. However, 48% of the total zooplankton biomass larvae held in the laboratory at 25 to during the period of larval residence. 40 ppt metamorphosed completely with They speculated that this decrease may no abnormalities (Reintjes and Pacheco be partly explained by larval feeding. 1966; Hettler 1981); and larvae Durbin and Durbin (1975) suggested trapped in a natural cove at Beaufort, that Atlantic menhaden in coastal North Carolina, transformed into waters may also alter the composition juveniles at 24 to 36 ppt (Kroger et of plankton assemblages by grazing on al. 1974).

certain size ranges.

Salinity affects temperature tolerance, activity, metabolism, and Important Atlantic menhaden growth. . Low salinities decreased predators include bluefish (Pomatomus survival at temperatures below 5 CC, saltatrix), striped bass (Morone and survival was poor at 6 'C in saxatilis), bluefin tuna (Thunnus freshwater (Lewis 1966). The effect thynnus), and sharks (Reintjes and of salinity on upper temperature Pacheco 1966). Atlantic menhaden tolerance was not significant (Lewis occurred in 13% of stomachs of sandbar and Hettler 1968). Larvae that sharks (Carcharhinus plumbeus). About Hettler (1976) reared at 5 to 10 ppt 46% of the menhaden were consumed whole exhibited significantly higher activi-and were 5-10 cm TL. Menhaden consumed ty levels, metabolic rates, and growth in more than one piece or partially rates than those reared at 28 to 34 consumed were 5-17 cm TL. This prey ppt. Lewis (1966) also noted slower was the second most frequently consumed growth at high salinities. Subtle type for sandbar sharks (Medved et al. physiological adaptations to low 1984). Because Atlantic menhaden are salinity may be an evolutionary eaten by predators in several eco- response to larvae "seeking" the systems, they are a direct pelagic link food-rich estuarine environment.

in the food web between detritus and Rogers et al. (1984) noted that pre-plankton and top predators. juveniles of many fishes, including 13

those of Brevoortia species, entered ted in the North Atlantic Region were estuarine habitats during seasonal between 10 and 13 'C (Ferraro 1980a).

peaks of freshwater influx when the The temperature range for the Middle area of low-salinity and fresh tidal Atlantic Region was 0 to 25 'C, but water was greatest. most eggs and larvae were collected at 16 to 19 'C (Kendall and Reintjes An important management consider- 1975).

ation is that, during the evolution of the Atlantic menhaden, estuarine zones The limits of larval temperature received freshwater from contiguous tolerance are also affected by accli-wetlands and riverine systems. How- mation time. Survival above 30 °C ever, channelization, diking of river (Lewis and Hettler 1968) and below 5 courses, ditching and draining of 'C (Lewis 1965) was progressively marginal wetlands, and urbanization extended by acclimation temperatures have reduced the freshwater retention closer to test values, suggesting that capacities of coastal wetlands. rapid changes to extreme temperatures Furthermore, extensive filling of are more likely to be lethal than estuarine marshlands has diminished prolonged exposure to slowly changing the area receiving runoff in many values. Winter shutdown of power plant locations. In combination, these operations may result in rapid changes cause rapid discharge of high temperature decreases near the effluent volumes of freshwater during brief discharge area. Mortality of juvenile periods and reduced amounts of fresh- Atlantic menhaden to a temperature water at other times. High inflows, decrease of 10 °C (from 15 to 5 C) was particularly those that occur in early less at rates of decrease of 6.7 °C/h spring after the arrival of pre- or lower than at faster rates. Winter juvenile menhaden, can expose fish to menhaden kills can be minimized by extreme fluctuations of temperature, reducing the rate of decrease as the turbidity, and other environmental power plant discharge is shut down conditions. Although the effects of (Burton et al. 1979).

altered freshwater flow regimes on Atlantic menhaden are not known, Hettler and Colby (1979) demon-effects on other estuarine-dependent, strated that photoperiod at least offshore-spawned fishes range from partly explains variation in resis-disappearance (Rogers et al. 1984) to tance to heat stress. Median lethal death (Nordlie et al. 1982). These time increased linearly with photo-effects are also mediated by tempera- period at 34 *C. They also speculated ture (Nordlie 1976). that it may be important to other types of physiological stress. Lewis Salinities of 10 to 30 ppt did and Hettler (1968) observed increased not affect developing embryos, though survival of juveniles at 35.5 'C with temperature did (Ferraro 1980a). increased dissolved oxygen (DO)

Mortality of embryos was complete at saturation. Burton et al. (1980) temperatures less than 7 °C and was reported a mean lethal DO concentra-significantly higher at 10 'C than at tion of 0.4 mg/l , but warned against 15, 20, and 25 °C. Time to hatching interpretation of this value as was significantly shorter at each "safe," in view of the interactive progressively higher temperature. nature of environmental factors.

Surface temperature in the spawning Westman and Nigrelli (1955) observed areas of the South Atlantic Region mass mortalities from gas embolism during the months of highest egg only in areas with highly variable capture were generally 12 to 20 'C salinity and organic pollution (Walford and Wicklund 1968). The sufficiently severe to make shellfish lowest temperatures at which Atlantic unfit for human consumption. Lewis menhaden eggs and larvae were collec- and Hettler (1968) observed decreased 14

survival at high temperatures by fish lack of significant differences affected by gill parasites. The between areas within years, although interaction of environmental factors this may have been due to the sampling must be considered when one defines regime. They speculated that PCB healthy ranges for-an organism. levels have remained somewhat high because of leakage from sources established prior to regulation and Substrate and System Features continued allowance of limited specialty uses. Menhaden oil products The association of the Atlantic carry the highest concentrations of menhaden with estuarine and nearshore such non-polar compounds and some systems during all phases of its life samples contained levels in excess of cycle is well documerted. It is evi- United States Food and Drug Admini-dent that young menhaden require these stration temporary tolerances, as of food-rich waters to survive and grow, 1977. Warlen et al. (1977) demonstra-and the fishery is concentrated near ted that 1"4C-DDT uptake by Atlantic major estuarine systems (June 1961a). menhaden is dose-dependent, with an Filling of estuarine wetlands, in assimilation value between 17% and addition to exacerbating extremes in 27%. Application of their model to .

environmental conditions, has physi- field data suggested that uptake was cally limited the nursery habitat by way of plankton and detritus.

available to Atlantic menhaden and Little information exists about the other estuarine-dependent species. toxicity of contaminants to Atlantic The relative importance, however, of menhaden.

different habitat types (i.e., sounds, channels, marshes) and salinity regimes has received little detailed Other Factors attention.

The seasonal depth distribution of Atlantic menhaden is tied to migra-Environmental Contaminants tion patterns. Some fish occur year-round in depths of 1 to 200 m (3 to In a study of chlorinated hydro- 656 ft). The role of turbidity in carbon residues in menhaden fishery Atlantic menhaden biology apparently products from the Atlantic and Gulf of has not been studied. Blaber and Mexico, Stout et al. (1981) showed Blaber (1980) proposed that gradients that overall levels have decreased of turbidity, nutrients, and salinity since the late 1960's, although signi- could provide cues that enable fry to ficant differences between years for locate estuarine nursery areas along levels of polychlorinated biphenyls the Australian coast. The "seeking" (PCB's) in the South Atlantic Region of turbid zones is probably related to and for dieldrin in the Middle differential mortality linked to food Atlantic Region could not be demon- supply and predation (Blaber and strated. There was also a general Blaber 1980; Norcross and Shaw 1984).

15

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19

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20

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21

Nicholson, W. R., and J. R. Higham, Reintjes, J. W. 1968. Development and Jr. 1965. Age and size composition oceanic distribution of larval of the menhaden catch along the menhaden. U.S. Fish Wildl. Serv.

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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 1989 Coastal Fishes and Invertebrates (Mid-Atlantic)--Atlantic Menhaden L fL PJionn~Q*n~ C'gecI, a.*iII en., Nl.I

7. A ihas S. Gordon Rogers and Michael J. Van Den Avyle Georgia Cooperative Fish and Wildlife Research Unit _

School of Forest ResourcesI C-'*,,,*,c*) ,CGNo.

University of Georgia Athens, GA 30602 IL Zeo..o..n 0C...,i.fo s PA..... 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 Station U.S Department of the Interior P.O. Box 631 Washington, 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 assist with environmental impact assessment. The Atlantic menhaden (Brevoortia tyrannus) is an important commercial fish along the Atlantic coast. In the South Atlantic Region, Atlantic menhaden spawn during winter in continental shelf waters. Adults then move inshore and northward 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 months at sea. Young fish move into the shallow regions of estuaries and seem to prefer vegetated marsh habitats. Atlantic menhaden are size-selective plankton feeders as larvae, and filter feeders as juveniles and adults. Due to their large population size, individual growth rates, and seasonal movements, Atlantic menhaden annually consume and redistribute large amounts of energy and materials. They are also important prey for large game fishes such as bluefish (Pomatomus saltatrix), striped bass (Morone saxatilis), and bluefin tuna (Thunnus thynnus). The Atlantic menhaden is associated with estuarine and nearshore systems during all phases of its life cycle. Young menhaden require these food-rich habitats to survive and grow. Destruction of estuarine wetlands has decreased nursery habitat available to Atlantic menhaden and other estuarine-dependent species.

Estuaries Feeding habits Fishes Growth Atlantic menhaden Fisheries Brevoortia tyrannus Habi tat Spawning IL A-landiy SI4.**,.. IS. 'O.ln f C:,I *'ThIR* l 21t NO. Oft UbUnclassified 23 UnlimiteduDistribution S,- .',*s at" '

Urn. .,. ÷:%-!*e

As the Nation's principal conservation agency, the Department of the Interior has responsibility for most of our nationally owned public lands and natural resources. This includes fostering the wisest use of our land and water resources, protecting our fish and wildlife, preserving the environmental and cultural values of our national parks and historical places, and providing for the enjoy-ment of life through outdoor recreation. The Department assesses our energy and mineral resources and works to assure that their development is in the best interests of all our people. The Depart-ment also has a major responsibility for American Indian reservation communities and for people who live in island territories under U.S.

administration.

U.S. DEPARTMENT OF THE INTERIOR 1 411(Vk1 1 AF FISH AND WILDLIFE SERVICE 1~]i TAKE PRIDE in America UNITED STATES DEPARTMENT OF THE INTERIOR POffAGE AND FEES PAID FISH AND WILDLIFE SERVICE UAL. DEPARTMENT OF ThE INTERIOR National Wetlands Research Center IMT-423 NASA-Slidell Computer Complex 1010 Gause Boulevard Slidell, LA 70458 OFFICIAL BUSINESS PENALTY FOR PRIVATE USE. $300