Library

National Wetlands Research Center

U. S. Fish and Wildlife Service

Biological Report 82(11.46)

April 1886

Lafayette, La. 70506 TR EL-82-4

Species Profiles: Life Histories and

Environmental Requirements of Coastal Fishes

and Invertebrates (South Atlantic)

AMERICAN SHAD

QL

155

.S63

no. 82

11.45

Fish and Wildlife Service

U.S. Department of the Interior

Coastal Ecology Group

Waterways Experiment Station

U.S. Army Corps of Engineers

Biological Report 82(11.45)

TR EL-82-4

April 1986

Species Profiles: Life Histories and Environnental Requirements

of Coastal Fishes and Invertebrates (South Atlantic)

AMERICAN SHAD

Douglas E. Facey

and

Michael J. Van Den Avyle

Georgia Cooperative Fishery Research Unit

School of Forest Resources

University of Georgia

Athens, GA 30602

Project Officer

John Parsons

National Coastal Ecosystems Team

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

National Coastal Ecosystems Team

Division of Biological Services

Research and Development

Fish and Wildlife Service

U. S. Department of the Interior

Washington, DC 20240
This series should be referenced as follows:

U.S. Fish and Wildlife Service. 1983-19 Species profiles: life histories

and environmental requirements of coast;ir'fishes and invertebrates. U.S. Fish

Wildl. Serv. Biol. Rep. 82(11). U.S. Army Corps of Engineers, TR EL-82-4.

This profile should be cited as follows:

Facey, D. E., and M. J. Van Den Avyle. 1986. Species profiles: life histories

and environmental requirements of coastal fishes and invertebrates (South

Atlantic) -- American shad. U.S. Fish Wildl. Serv. Biol. Rep. 82(11.45).

U.S. Army Corps of Engineers, TR EL-82-4. 18 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 Coastal Ecosystems Team

U.S. Fish and Wildlife Service

NASA-Slide11 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
Multiply !Y

millimeters (mn) 0.03937

centimeters (cm) 0.3937

meters (m) 3.281

kilometers (km) 0.6214

square meters (m2)

square kilometers (bn2)

10.76

0.3861

2.471

To Obtain

Inches

Inches

feet

miles

hectares (ha)

square feet

square miles

acres

liters (1)

cubic meters (m3)

cubic meters

0.2642

35.31

0.0008110

gallons

cubic feet

acre-feet

milligrams (mg)

grams (9)

kilograms (k

metric tons 9

)

t)

metric tons

kilocalories (kcal)

0.00003527

0.03527

2.205

2205.0

1.102

3.968

ounces

ounces

pounds

pounds

short tons

British thermal units

Celsius degrees 1.8(Y) + 32 Fahrenheit degrees

CONVERSION TABLE

Metric to U.S. Customary

inches 25.40 millimeters

inches 2.54 centimeters

feet (ft.) 0.3048 meters

fathoms 1.829 meters

miles (mi) 1.609 kilometers

nautical miles (rmi) 1.852 kilometers

square feet (ft2)

acres

square miles (mi2)

0.0929 square meters

0.4047 hectares

2.590 square kilometers

gallons (gal) 3.785 liters

cubic feet (ft3) 0.02831 cubic meters

acre-feet 1233.0 cubic meters

ounces (02)

pounds (lb)

short tons (ton)

28.35

0.4536

0.9072

British thermal units (Btu) 0.2520

Fahrenheit degrees 0.5556("F - 32)

grams

kilograms

metric tons

kilocalories

Celsius degrees

U.S. Customary to Metric

iv
CONTENTS

'PREFACE .................................. iii

CONVERSION TABLE ............................. iV

ACKNOWLEDGMENTS .............................. vi

NOMENCLATURE/TAXONOMY/RANGE . . . .

MORPHOLOGY AND IDENTIFICATION AIDS

REASON FOR INCLUSION IN THIS SERIES

LIFE HISTORY . . . . . . . . . .

Adult Migration and Spawning . .

Eggs and Larvae . . . . . . . . .

Juveniles and Adults . . . . . .

GROWTH CHARACTERISTICS

COMMERCIAL AND SPORT FISHERIES' : :

Stock Abundance . . . . . . . . .

Gear . . . . . . . . .

ManagemeAt' : : . . . . . . . . .

The Sport Fishery . . . . . . . .

Population Dynamics . . . . . . .

ECOLOGY . . . . . . . . . . . . . .

Feeding . . . . . . . . . . . . .

Potential Competitors . . . . . .

Predators and Diseases . . . . .

ENVIRONMENTAL REQUIREMENTS . . . .

Temperature . . . . . . . . . . .

Salinity . . . . . . . . . . . .

Dissolved Oxygen . . . . . . . .

Water Movement

Turbidity, Substiate; And Depth :

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LITERATURE CITED . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Page

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6

7

7

9

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10

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::

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13

13

14

15

V
ACKNOWLEDGMENTS

We thank Ronald J. Essig, Georgia Department of Natural Resources, Brunswick; Nan C. Jenkins, South Carolina Wildlife and Marine Resources Department,

Charleston; and John W. McCord, South 'Carolina Wildlife and Marine Resources

Department, Charleston, for reviewing the manuscript. J. Ernest Snell, Supervisory Statistician, National Marine Fisheries Service, Brunswick, Georgia,

provided recent data on American shad harvest. Jennifer Biggers prepared Figure

1.

Vi
Figure 1. American shad.

AMERICAN SHAD

NOMENCLATURE/TAXONOMY/RANGE

Scientific name . . . . . . . . Alosa

sapidissima (Wilson)

Preferred comnon name . . . . American

shad (Figure 1)

Other common names . . . . shad, white

shad, comnon shad, Atlantic shad,

North River shad, Potomac shad,

Connecticut River shad, Delaware

shad, Susquehanna shad, alose (Scott

and Crossman 1973).

Class . . . . . . . . . . Osteichthyes

Order . . . . . . . . . . Clupeiformes

Family . . . . . . . . . . . Clupeidae

Geographic range: The American shad

is native to the east coast of

North America from northern

Florida to Newfoundland and is

most abundant from North Carolina

to Connecticut (Figure 2). The

species is anadromous and lives in

fresh-, brackish-, and saltwater

during its various life stages.

American shad were successfully

introduced into the Sacramento and

Columbia Rivers on the Pacific

coast in the late 1800's, and now

are established from southern

California northward to Cook

Inlet, Alaska, and the Kamchatka

Peninsula on the Asiatic side of

the North Pacific. Attempts to

introduce American shad into Lake

Ontario, the Mississippi River

drainage, peninsular Florida, and

Great Salt Lake have not been

1
NORTH CAROLINA B,-

\

-- “Y

_--

SOUTH CAROLINA’b_

MILES

5 -

iEORGl3 ATLANTIC OCEAN

NOTE: Dlotrlbutlon holudoo all

l rtuarlor end nmod rivorr.

2 K\ KILOMETERS

Figure 2.

Region.

Major spawning rivers of the American shad i n the South

2
successful (Scott and Crossman

1973). Millerton Lake, a reservoir on the San Joaquin River

in California, supports the

only known reproducing landlocked

population of American shad

(Lambert et al. 1980).

MORPHOLOGY AND IDENTIFICATION AIDS

The following description of

American shad is summarized from

Hildebrand (1963) and Scott and Crossman (1973).

The American shad is the largest

member of the herring family (Clupeidae). Females may reach a total

length (TL) of 600 mm and a weight of

5.4 kg. The body is slender and

strongly compressed; body depth is

25-38% of standard length (SL) and

increases with age. The head is

broadly triangular; the gill membranes

are entirely free from the isthmus.

The maxillary is broad and extends to

the middle of the eye in the young,

and generally to below the posterior

margin of the eye in adults. The gill

rakers increase ’ number

proportionate 1ength':ith age;

and

adults

usually have 59-73 rakers on the

lower limb of the first gill arch.

Teeth on the jaws and tongue of young

shad are lacking in adults. The

American shad has one dorsal fin

(15-19 rays) of moderate height, a

deeply forked caudal fin, and an anal

fin with 18-24 rays. There are 35-38

well-developed ventral scutes, 19-23

in front of the pelvic fins and 12-19

behind. American shad have 53-59

(usually 55-58) vertebrae.

Live specimens have a greenishto bluish-metallic luster on the back

and are bright silver on the sides. A

dark spot on the shoulder, just behind

the posterior edge of the operculum,

is usually followed by 3 to 27 smaller

spots or dots. There is sometimes a

second row of 1 to 16 spots below the

first, and rarely a third row of 2 to

9 spots below the second.

The American shad lacks the

greatly elongated last dorsal fin ray

present in both the gizzard shad

(Dorosoma cepedianum) and the threadfm(D. petenense). It has more

than 55 gin rakers on the lower limb

of the first gill arch and usually has

at least four black spots in a horizontal row behind the ooerculum. The

congeneric alewife (A.

and blueback herring

have fewer than 55 air1 rakers and

only one prominent bla:k spot near the

upper rear edge of the operculum. The

hickory shad (A. mediocris) has 18-23

gill rakers on-the lower limb of the

first gill arch and it has teeth in

the lower jaw at all ages (Dahlberg

1976). In adult American shad, the

maxillary extends at least to the

posterior margin of the eye, whereas

in the alewife and blueback herring it

extends only to the midpoint of the

eye. Alosa species have a distinct

median notch in the upper jaw which

separates them from other Clupeid

species.

REASON FOR INCLUSION IN THIS SERIES

The American shad has supported

major commercial fisheries along the

Atlantic coast since the early 1800's

and was the most valuable food fish

along the east coast before World War

II (Rulifson et al. 1982). Commercial

landings have declined sharply since

the early 1900's; they exceeded 50

million pounds in 1896, but fell to

about 2 million pounds in 1976. In

many Atlantic coast rivers that still

support runs, the sport fishery for

American shad has become more important than the commercial fishery.

The decline of American shad has

been attributed to dams, overfishing,

habitat destruction, and pollution

(Sholar 1977; Rulifson et al. 1982).

Different factors have been blamed in

different river systems and in some

instances the combined effects of

several factors may be responsible;

however, data are too scarce to
satisfactorily explain the drastic

reduction in the shad population.

Because the American shad is an

anadromous fish, it is susceptible

to perturbations of freshwater,

estuarine, and marine habitats, and

the needs of the species should be

considered in riverine and coastal

development projects.

LIFE HISTORY

The life history of American shad

on the Atlantic coast was summarized

by Walburg and Nichols (1967). Ulrich

et al. (1979a) and Rulifson et al.

(1982) provided updated life history

reviews for the species in the southeastern United States. Much of the

information in this section was taken

from these two references.

Adult Migration and Spawning

The following offshore migratory

patterns for the adult American shad

were proposed by Neves and Depres

(1979) and are based on field surveys

and an extensive review of the

literature. In the spring after

spawning, adult shad probably migrate

to the Gulf of Maine or to an area

south of Nantucket Shoals (off

southern New England) where they

remain through the Sumner and early

fall. These offshore waters are 50 to

100 m deep, and the water temperatures range from 3O to 15' C. In

fall and winter, schools of adult

fish gradually move southward and

overwinter offshore between Long

Island and Nantucket Shoals. They also

tend to congregate in Middle Atlantic

coastal waters during late winter and

spring. Shad that were spawned near

the northern range of their distributiot, move northward along the coast

as waters warm above 3" C. The south

Atlantic populations migrate southward

along the coast (within the 15' C

isotherm), thereby reaching their home

rivers by late winter or early spring.

Most American shad return to

their natal streams to spawn.

Significant differences in morphological characters among shad from

different Atlantic coastal rivers

support the belief that there are

discrete spaming populations (see

Walburg and Nichols 1967). Although

this homing tendency is strong, some

individuals ascend other rivers to

reproduce. The homing behavior of the

shad in the Connecticut River may be

related to olfaction and orientation

to the river current (Dodson and

Leggett 1974). These mechanisms

probably hold for other shad

populations as well.

Water temperature changes are

partly responsible for the timing of

the spawning migration; most shad

enter rivers when water temperatures

are betwen 10" and 15" C (Leggett and

Whitney 1972). In the St. Johns

River, Florida, the migration usually

peaks in January when the water

temperature is about 15' C (Ulrich et

al. 1979a). In general, the peak of

the spawning migration becomes progressively later in the year from

south to north.

Along the South Atlantic Bight

American shad begin their spawning

migration in rivers during winter and

early spring. They begin to move up

the St. Johns River, Florida, as early

as November, peaking from mid-January

to mid-February (Ulrich et al. 1979a).

The spawning run begins in early

January in Georgia and South Carolina

and in January and February in North

Carolina (Ulrich et al. 1979a; Sholar

1977). Some spawning runs may

continue as late as April or May. In

eastern Canada, some shad do not enter

rivers until early June (Walburg and

Nichols 1967).

American shad usually spawn in

freshwater over substrates of sand,

gravel, and mud (Rulifson et al. 1982;

Stier, in press). Spawning probably

begins in late afternoon or evening

and continues until about midnight

4
(Ulrich et

spawning has

turbid rivers

American

al. 1979a). Diurnal

also been reported in

(Chittenden 1976a).

shad usually spawn at Eggs and Larvae

water temperatures of 14" to 21" C

(extremes 8" to 26' C) and require

dissolved oxygen concentrations of at

least 5.0 mg/l (Walburg and Nichols

1967).

The fecundity of American shad

varies among natal rivers and the age

of the fish. The range is about

100,000 to 600,000 eggs per female

(Cheek 1968). Leggett and Carscadden

(1978) showed that fecundities along

the east coast decreased from south to

north (Table 1). The fecundity of

shad offshore from North Carolina

ranged from 197,000 to 457,000 eggs

per female and increased with size and

age.

Unfertilized American shad eggs

are pale amber or pink transparent

spheres about 1.27 mn in diameter

(Ulrich et al. 1979a). After fertilization and water hardening, the

diameters increase to about 2.5-3.8 mm

(Walburg and Nichols 1967). The eggs

are slightly heavier than water,

nonadhesive, and require water currents sufficient to buoy the eggs

during incubation. Shad eggs generally hatch in 4-6 days at 15"-18' C, but

the length of the incubation period

depends on water temperature (see

ENVIRONMENTAL REQUIREMENTS section for

further details).

Most American shad from rivers in

the southeastern United States die

after they spawn, but the occurrence

of repeat spawners may exceed 60% in

northeastern rivers (Table 2). Reported percentages of repeat spawners are

near zero for rivers south of the

Neuse River, North Carolina (Table 2).

More recent evidence, however, indicates that these rivers may have a few

repeat spawners (J. W. McCord, South

Carolina Wildlife and Marine Resources

Department, Charleston; personal

communication). Adults that survive

spawning move downriver and migrate to

The larvae of American shad are

about 7-10 mm long when they hatch

(Walburg and Nichols 1967). On the

second or third day after hatching,

small teeth appear on the lower jaw

and in the pharynx, and the yolk is

usually absorbed b

J

the fifth day

(Ulrich et al. 1979a . Feeding begins

between the 10th and 12th days, and

growth is fairly rapid. Larvae

develop into juveniles after 4-5 weeks

when they are about 25 mm long

(Walburg and Nichols 1967).

Table 1. Mean virgin (first spawn) and lifetime fecundities of American shad

from five Atlantic coast rivers (Leggett and Carscadden 1978).

the Gulf of Maine (Walburg and Nichols

1967).

Location

Thousands of egqs

Virgin Lifetime

Miramichi River, New Brunswick 129 258

St. John River, New Brunswick 135 273

Connecticut River, Connecticut 256 384

York River, Virginia 259 327

St. Johns River, Florida 406 406

5
Table 2. Percent of American shad that spawn more than once in Atlantic coast

rivers.

Location Source

St. John River, New Brunswick

Connecticut River, Connecticut

Hudson River, New York

Susquehanna River, Maryland

James River, Virginia

Chowan River, North Carolina

Neuse River, North Carolina

Cape Fear River, North Carolina

Edisto River, South Carolina

Ogeechee River, Georgia

St. Johns River, Florida

57

37

27

15

7

<l

:

0

Leggett

Leggett

Leggett

Leggett

Leggett

Holland

Hawkins

Fischer

Leggett

Leggett

Leggett

and Carscadden (1978)

and Carscadden (1978)

and Carscadden (1978)

and Carscadden (1978)

and Carscadden (1978)

q;;8;ylverton (1973)

(1980)

and Carscadden (1978)

and Carscadden (1978)

and Carscadden (1978)

Juveniles and Adults

Juvenile American shad usually

form schools and gradually move downriver at a rate dictated by water

temperature and current velocity

(Williams and Bruger 1972; Stier, in

press). Downstream portions of

rivers, tidally influenced freshwater

zones, and estuaries are used as

nursery areas (Rulifson et al. 1982).

Juveniles grow rapidly and commonly

exceed 90 mm total length when they

begin migrating to the sea (see Rulifson et al. 1982). Emigration from

rivers usually begins when !ater

temperatures

(Walburg and Nidcrhoo4s %$, b?i5juvef

niles in some northern populations may

remain in rivers and estuaries

throughout the first winter (Stier, in

press). In the sea, the juveniles

migrate to the Bay of Fundy and then

to the Gulf of Maine where they join

the adults each summer (Dadswell et

al. 1983). They probably move southward and spend the winter in the midAtlantic area (Neves and Depres 1979).

American shad from North Carolina

become sexually mature when they are 3

to 5 years old (Sholar 1977). Catch

data from southern rivers, in which

repeat spawners are rare, show that

most spawning males are 3-5 years old

and most females are 4-6 years old

(Loesch et al. 1977; Fischer 1980;

Hawkins 1980; Rulifson et al. 1982).

GROWTH CHARACTERISTICS

American shad that spawn in

southeastern rivers usually are 3-6

years old and reach 400-525 mm fork

length. Because few fish survive to

spawn again, their life span is

shorter than that of shad in northern

populations. The age of American shad

can be determined from scales, which

show growth rings as well as marks

that correspond to spawning migrations

(Judy 1961). The average size at a

given age is fairly consistent among

shad from different rivers in the

South Atlantic (Table 3). Females

tend to be larger than males of the

same Holland and Yelverton

(1973)aFedicted age (A) from fork

length (FL, cm) for American shad

(sexes combined) from offshore North

Carolina as:

A = 33.86 FL"*17.

6
Table 3. Age and mean fork length (mm) of American shad caught in rivers of the

South Atlantic.

Age group

Location Sex I II III IV v VI VII VIII Source

Pamlico Sound

and River,

North Carolina

Neuse River,

North Carolina

Cape Fear River,

North Carolina

Santee River,

South Carolina

Savannah River,

South Carolina/

Georgia

St. Johns River,

Florida

M 404 425 452

F 429 464 486 523

M 375 401 418 435 455

F 428 460 482 502

M 375 415 439 463

F 425 466 491 518

M 370 415 425 470

F 440 470 500 525

M 405 445 465

F 440 475 505 525

M 175 277 345 394

F 180 284 353 409 447

Hawkins

(1980)

547

Hawkins

(1980)

Fischer

(1980)

Ulri ch et

al. (1983)

Ulri ch et

al. (1983)

LaPoin e

(1958) dr

aAs cited by Holland and Yelverton (1973).

They also predicted body weight (W,

kg) from FL (cm) as:

W = (6.84 x 10-6) FL3*23.

Fork length (cm)-weight (g) regression

equations for fish taken from

different areas in South Carolina were

W = 88.29 FL - 2509.31 for males and W

= 111.87 FL - 3390.58 for females (N.

C. Jenkins, South Carolina Wildlife

and Marine Resources Department,

Charleston; personal communication).

Regressions by sex for specific river

systems in South Carolina were

published by Ulrich et al. (1979b).

Walburg (1956) found that juvenile

shad from six rivers along the

Atlantic coast from Florida to

Connecticut were similar in size

during late summer, even though they

spawned earlier in the south than in

the north.

COMMERCIAL AND SPORT FISHERIES

Stock Abundance

Historically, the Atlantic coast

fishery for American shad has been

one of the most important anadromous

fisheries in North America (Rulifson

et al. 1982). By the mid 1700's, shad

supported profitable fisheries from

the Chesapeake Bay to Maine. In the

early 1800’s, access to northern

markets was improved by advances i n

transportation, and shad fisheries

became important as far south as

Florida. Shad increased in economic

importance during the 1800's and

7
supported fisheries in every State

along the Atlantic coast. Since the

record-setting catch of more than 50

million pounds in 1896, annual landings generally declined. The reported

catch for 1976 was about 2 million

pounds (NMFs 1980).

Commercial fisheries for shad

in southeastern coastal rivers have

shown the same historical declines.

The North Carolina shad harvest peaked

near 9 million pounds in 1896, but

declined to less than 200,000 pounds

in 1980 (Table 4). The catch in

Georgia exceeded 1.3 million pounds in

1908, but was less than 200,000 pounds

in 1981. The trend has been the same

in South Carolina and Florida.

Increased landings after 1977 (see

Table 4) may have been partly due to

collection of more accurate catch data

(e.g., Ulrich et al. 1983).

A survey of licensed shad fishermen in South Carolina showed that less

than 5% relied on commercial fishing

to make a living , and that even these

fishermen derived less than 10% of

their annual income from shad fishing

(Ulrich et al. 1979a). Most of those

surveyed fished only for recreation

and home consumption. Among Georgia

shad fishermen surveyed during 1980-

1982, about 12% indicated that they

derived at least 50% of their income

Table 4. Cormnercial landings (thousands of pounds) and dockside value (thousands

of dollars) of American shad in the South Atlantic States, 1960-1962 (Ulrich et

al. 1979a; Hawkins 1980; Rulifson et al. 1982; Essig 1983; unpublished data,

Georgia Department of Natural Resources and U.S. National Marine Fisheries

Service).

North Carolina South Carolina Florida

Year Landings Value Landings Value LanPings Value LandingsValue

1960 507

1961 673

1962 765

1963 693

1964 640

1965 1069

1966 701

1967 777

1968 842

1969 719

1970 953

1971 680

1972 468

1973 321

1974 369

1975 241

1976 167

1977 121

1978 402

1979 278

1980 199

1981 352

1982 412

127

168

191

168

127

214

170

1;8

137

193

117

112

1::

83

65

1::

122

88

190

183

106 34

1105 :;

120

120 %i

176 61

119 34

132

110 ;1

17748 35;

1:; t k!

;: :;

62 37

32 20

2:; 197 54

197 155

271 215

446 283

243 198

533 176

404

527

331

314

376

386

334

569

618

532

420

344

239

162

182

93

118 84

238 113

268

188

196

198

88

100

127

100

139

182

140

133

112

91

63

99

57

172

148

166

468

425

760

590

613

758

319

531

390

218

253

120

1::

1:

1::

115

181

241

181

27

24

9

8
income from some form of commercial

fishing (Essig 1983).

The major shad-producing rivers

in North Carolina are the Tar-Pamlico,

the Cape Fear, the Northeast Cape

Fear, the Chowan, and the Neuse. At

one time, the Neuse River was considered to be the most important shad

stream between the St. Johns River,

Florida, and the James River, Virginia

(Walburg and Nichols 1967; Rulifson et

al. 1982). The principal shad producing area in South Carolina is Winyah

Bay and its major tributaries, the

Waccamaw and Pee Dee Rivers (Ulrich et

al. 1979a). Other waters supporting

shad fisheries are the Santee River,

Charleston Harbor (and its tributaries, the Cooper and Ashley Rivers),

the Edisto River, and the Combahee,

Ashepoo, and Coosawhatchie Rivers.

The Savannah River supports shad runs

that benefit fishermen from both South

Carolina and Georgia. Besides the

Savannah River, Georgia shad fishermen

use the Ogeechee, Altamaha, Satilla,

and St. Marys Rivers. The Altamaha

now supports Georgia's largest shad

fishery (Michaels 1980), followed

closely by the Savannah River fishery

(Essig 1983). Most shad fishing in

Florida is in the St. Johns River.

Gear

From about 1960 to 1970, North

Carolina shad fishermen primarily used

drift and anchored gill nets (56%),

pound nets (41%), and haul seines (3%)

(Sholar 1977). The predominant gear

at the turn of the century was pound

nets. Gill nets have recently become

the major gear because of the

increased efficiency of nylon monofilament twine and low cost compared to

pound nets and seines (Sholar 1977).

Shad fishermen in South Carolina

have used a variety of fishing gear

over the years. Gill nets have been

used since the late 1800's. Seines,

dip nets, cast nets, and fyke nets

have been popular with some fishermen

at one time or another, but drift and

set gill nets are now the only gear

fished commercially (Ulrich et al.

1979a).

Gill nets also have been the

preferred gear of shad fishermen in

Georgia since the late 1800's; bow

nets, pound nets, and fyke nets also

have been used. Since the 1960's,

commercial shad fishing in Georgia has

been primarily accomplished with gill

nets; drift gill nets are preferred

over set gill nets. Drift gill nets

accounted for 82% of the Georgia shad

landings in 1979-1982 (Essig 1983).

Some fishermen still use hand lines

and otter trawls.

Commercial shad fishing in Florida

is restricted to gill netting and

angling (Capt. L. Shelfer, Florida

Marine Patrol, personal communication).

Manaqement

In North Carolina, fishing for

American shad is heaviest in January

through April; there are no closed

seasons (Rulifson et al. 1982). The

open season for shad fishing in South

Carolina varies somewhat among the

rivers, but generally fishin is legal

from February 1 to April 30 9Ulrich et

al. 1979a). The season in Georgia is

closed from May 1 through December 31.

The open season is established each

year by the Commissioner of Natural

Resources, within the period January 1

- April 30 (R. J. Essig, Georgia

Department of Natural Resources,

personal communication). Florida

waters are closed to shad fishing

between March 15 and November 15, and

the open season generally extends from

!II;;;E;r 31 to March 1 (Capt. L.

Florida Marine

personai communication).

Patrol;

The minimum legal mesh size for

comnerical gill netting of American

shad in South Atlantic States is

between 11.4 and 14.0 cm (stretch

mesh). Regulations controlling the

number and length of nets and legal

9
fishing hours and days vary within and

among river systems.

The Sport Fishery

Sport fishing for American shad

with hook and line is common in the

Cape Fear and Tar Rivers in North

Carolina (Sholar 1977). In late

winter, drift gill netting for American shad, hickory shad, and blueback

herring is a popular sport in the

Neuse River, North Carolina (Hawkins

1980). Sport fishermen in South

Carolina and Georgia often fish for

American shad with small artificial

lures such as shad darts or small

spoons (Ulrich et al. 1979a). Bow

nets also are used for sport fishing

in South Carolina and Georgia. The

American shad has become a popular

sport fish in some Florida rivers,

particularly the St. Johns (Williams

et al. 1975).

Population Dynamics

Selective fishing for females and

general overexploitation are problems

rivers

~~lanmtai"cY States.

along the South

Mature female shad

( "roe" shad) are worth about twice as

much per pound as males ("buck" shad)

because the eggs are highly sought as

{;cK& by consumers (Ulrich et al.

Fishermen typically direct

their-fishing toward the more valuable

roe shad, which could reduce egg production and potential recruitment.

This selectivity for females also

biases estimates of population sex

ratios that are based on commercial

catch statistics. Samples of shad

taken by haul seine in the Northeast

Cape Fear River indicated

male:female ratio of 4.7:1. Shola:

(1977) suggested that the fishery may

be overexploiting the females.

Overexploitation of females could

seriously affect recruitment in future

years. In the Connecticut River, 64%

of the annual variation in juvenile

shad production was related to the

number of adults reaching the spawning

grounds (Marcy 1976). Leggett (1976)

related the number of adults that

reached spawning grounds in the Connecticut River with recruitment in the

next year as:

R = Ne0.7118( l-N/86.59)

,

where N was the number of eggs produced by the parent stock, and R was

recruitment.

Shad harvest in the southeastern

states generally includes males that

are 4-5 years old and females 5-6

years old (Table 5). However, in the

Altamaha River, Georgia, and the St.

Johns River, Florida, 3-year-old males

contribute a major share of the catch.

Rates of annual fishing mortality for

American shad vary from 15% to 66% in

southeastern rivers (Table 6).

ECOLOGY

Feeding

American shad consume a variety

of invertebrate organisms; small

fishes are an important part of the

diet in some areas. Shad larvae (14-

28 mm long) in the Connecticut River

above Holyoke Dam, Massachusetts,

mainly consumed cyclopoid copepods,

midge larvae, midge pupae, and Da hnia

*il;;ev;;quethean;am;eed

area

*

crustacean zooplankton, midge larvae

and pupae, caddis fly larvae, and

adult insects. These findings led

Levesque and Reed (1972) to conclude

that juvenile shad probably are

opportunistic feeders, although they

select most of their food from the

water column, rather than from the

bottom or near the surface.

The stomach contents (volume) of

juvenile shad in two tributaries of

the York River, Virginia, were 1%

amphipods, 28% aquatic insects, and

71% terrestrial insects (Massmann

1963). The mean volume of food per

individual shad was seven times

10
Table 5. Age composition (%) of American shad populations in rivers of the South

Atlantic Bight.

River system

and year

Age group

Sex II III IV v VI VII VIII Source

Albemarle Sound M 1 18 64

1978 F 4 56

Neuse River M 5 39 46

1977-1979 F 6 58

3: 1

1 1

Cape Fear River

1978-1979

Waccamaw-Pee Dee M <l 9 68

iii 4

Ulrich et al.

drainage 1982 F ~1 27 (1983)

Edisto River

1982 MF 134 834 4: 5

Ulrich et al.

(1983)

Savannah River M 7 80 Ulrich et al.

1982 F <l 46 :i 33 (1983)

Altamaha River M 1 3 49 36 <l

1967-1968 F Cl 26 62

1:

~1 <l

St. Johns River M 23

;: 1% lf <l

Williams et

1973 F cl 2 al. (1975)

MF 1 5918 4339 3: 3

Johnson et al.

(1978)

Hawkins (1980)

Fischer (1980)

Godwin (1968)

Table 6. Fishing mortality rates of American shad in southeastern rivers (from

Rulifson et al. 1982).

River system Year Mortality (%) Source

Neuse River, North Carolina

Waccamaw-Pee Dee

system, South Carolina

1957 65.0

1974 33.9

1975 29.0

1976 18.5

Edisto River, South Carolina 1955

Ogeechee River, Georgia

Altamaha River, Georgia

1954

1967

1968

St. Johns River, Florida 1960 15.0 Walburg (1960)

Walburg (1956)

Crochet et al.

(1976)

20.0

66.0

48.6

43.3

Walburg (1956)

Sykes (1956)

Godwin (1968)

11
greater at upriver stations, where

terrestrial insects were the predominant food, than at downriver

stations, where aquatic insects were

the principal food. Massmann (1963)

cautioned against estimating potential

fish production solely from data on

plankton and benthos because the

most important prey in his study came

from wooded areas bordering the river

and not from the river itself.

Walburg (1956) compared the food of

juvenile shad caught during August

from the St. Johns River, Florida,

Ogeechee River, Georgia, Neuse River,

North Carolina, Pamunkey River,

Virginia, Hudson River, New York, and

the Connecticut River, Connecticut.

Insects and crustaceans were the

primary food (by frequency of occurrence) in all rivers. Juvenile shad

also eat small fishes. Of 15 juveniles (87-141 mm long) caught off the

North Carolina coast, 12 had at least

one striped anchovy -(Anchoa he setus)

i n their stomachs (Hollan+-?iiid

Yelverton 1973). Juvenile shad (73-88

mm long) from the St. Johns River,

Florida, fed on the larvae of bay

anchovies (Anchoa mitchelli) and

mosquitofish (Gambusiaaffinis)

(Williams and arum). me

shad begin feeding in the late

afternoon; feeding intensity is

greatest during the early evening and

declines from midnight to midday

(Massmann 1963; Levesque and Reed

1972).

When adult American shad are offshore, they are believed to be planktivorous, consuming mainly copepods,

mysids, and other zooplankters (Hildebrand 1963; Vinogradov 1981; Rulifson

et al. 1982; Stier in press). The

stomachs of all 41 adults caught off

the coast of North Carolina contained zooplankton, including amphipods, copepods, isopods, cumaceans,

and larval decapods (Holland and

Yelverton 1973). Fish remains in

39 stomachs indicated that adult shad

are not exclusively planktivorous.

Vinogradov (1981) showed that the

types of zooplankton eaten by adult

shad in Nantucket Shoals varied with

the time of day. Adult shad usually

do not feed while migrating upstream

to spawn (Rulifson et al. 1982;

Chittenden 1976b).

Potential Competitors

Domermuth and Reed (1980) evaluated the potential for food competition among sympatric juvenile American

shad, juvenile blueback herrina. and

pumpkinseed sunfish (Lepomis

in the Connecticut River.

setts. They reported that prey selection or foraging locations differed

among the species. Pumpkinseeds ate

mainly benthic prey and were, therefore, not directly competing with the

clupeids, which fed mainly on plankton and drift organisms. Cladocerans

made up nearly 30% (by volume) of the

stomach contents of the shad and 47%

of the diet of the herring, but competition was probably light because

the two clupeids tended to consume

cladocerans from different families.

Copepods made up less than 2% of the

diet of shad but nearly 18% of the

diet of herring.

Chironomid larvae and pupae made

up 53.2% and 35.0% of the diet of shad

and herring, respectively, but since

chironomids were highly abundant in

the river, the sharing of this food

resource by shad and herring probably

had no adverse effect on either

species. Terrestrial adult insects

contributed 15% of the shad diet, but

were not present in the stomachs of

herring. Herring apparently fed only

in the water column, whereas shad fed

at the water's surface as well as in

the water column.

Predators and Diseases

Larval and juvenile American shad

probably fall prey to a variety of

predators. Young shad are eaten by

American eels (An uilla rostrata) and

striped bass ke saxatilis)

(Walburg and Nichols). Oncerlshad

enter the ocean, they are probably

17
eaten by a variety of offshore predators, including sharks, tuna, and

porpoises (Walburg and Nichols 1967).

Adult shad in rivers seem to have few

predators.

Shad are relatively free of

severe parasitic infestations. Nematodes, sea lice (isopods), roundworms,

trematodes, and acanthocephalans are

common parasites. Sea lampreys

(Petrom zon marinus) and freshwater

ld(IchFn spp.) have been

observed attac e to adult shad in the

Connecticut River (Walburg and Nichols

1967; Rulifson et al. 1982).

ENVIRONMENTAL REQUIREMENTS

Temperature

Water temperatures are critical

to American shad during their life

cycle. Most spawn at water temperatures between 14" and 21" C; extremes

&

were 8" and 26" C (Walburg and Nichols

1967). Leggett and Whitney (1972)

reported maximum survival of eggs and

larvae at X.5-26.6" C. Leim (1924)

reported that temperatures of 7" - 9'

C were lethal to eggs and larvae and

that temperatures of 20.0' - 23.4" C

caused extensive larval abnormalities.

The lower thermal tolerance limit of

juvenile shad has been reported as 2"-

4' C (Rulifson et al. 1982). Below 6"

C; juveniles lost equilibrium, ceased

feeding, and moved slowly (Chittenden

1972). Young shad can sense and avoid

potentially lethal temperatures (Moss

1970). Shad have been collected offshore at water temperatures from 3' to

15" C, but most live in a temperature

range from 5" to 13' C (Neves and

Depres 1979).

Salinity

Although American shad eggs are

deposited and hatch in freshwater,

some can tolerate moderate salinities.

Eggs were successfully hatched in

salinities ranging from 7.5 parts per

thousand (ppt) at 12" C to 15 ppt at

17" c. No eggs hatched at a salinity

of 22.5 ppt. Juvenile shad can tolerate sharp salinity changes, which

allows them to use both freshwater and

brackish waters as

(Chittenden 1973). Adu!$seL&e aiFzi

saltwater to freshwater during their

spawning migration, typically remaining in estuaries for 2-3 days. Adult

shad moved directly from saltwater to

freshwater started to die after 5

hours (Leggett and O'Boyle 1976).

Dissolved Oxygen

American shad usually spawn in

flowing water at dissolved oxygen (DO)

concentrations exceeding 5.0 mg/l

(Walburg and Nichols 1967). DO of at

least 4.0 mg/l is necessary in spawning areas (Chittenden 1973). Shad

eggs in the Neuse River, North

Carolina, were found in waters with

oxygen concentrations between 6 and 10

mg/l (Hawkins 1979). Mortality of

eggs and larvae exposed to DO concentrations of 2.5 to 2.9 mg/l was about

50% (Stier, in press), and mortality

of eggs was 100% at DO's below 1.0

mg/l (Carlson 1968). Larvae lost

equilibrium at a DO of 3.0 mg/l; many

died at DO's below 2.0 mg/l; and all

died at 0.6 mg/l (Chittenden 1969).

Juvenile shad seem to prefer high DO

concentrations when exposed to a

gradient but can probably survive low

DO (0.5 mg/l) for several minutes if

they have access to DO above 3.0 mg/l

(Dorfman and Westman 1970). Minimum

DO's of 2.5 - 3.0 mg/l are probably

sufficient to allow juvenile migration

through polluted waters (Chittenden

1973). Severely low DO concentrations

in rivers can prevent the passage of

adult shad to spawning

(Rulifson et al. 1982).

areas upstream

Water Movement

Proper development of shad eggs

requires water velocities that keep

the eggs suspended in the water

(Sholar 1977). Spawning is common in

water currents with a velocity from

30.5 to 91.4 cm/set (Walburg 1960).
Water velocity also affects energy

expenditures by adult shad that must

move upstream to reach spawning areas

(Glebe and Leggett 1981).

Dams block American shad migrations to upstream spawning areas, but

several methods can be used to pass or

lift the fish over dams. The earliest

designs often failed because water

depths and flows near fishways did not

attract adult shad. Most effective

fishways collect fish moving into one

or several entrances and direct them

to a single fish-passage facility.

Walburg and Nichols (1967) reviewed

the history and efficiency of shad

fishways along the Atlantic coast;

information in the remainder of this

section comes from their report.

Fishway collection systems for

shad should be at least 1.2 m deep

and 2.4 m wide. Water velocities

should not exceed 1 m/set. The

entrances to the collection system

should be at least 1.8 m wide and have

a depth of at least 0.3 m, preferably

1.0 m. The velocity at the entrance

should be 1.5-2.1 m/set.

After shad have been attracted

into the collection system, several

methods can be used to pass or lift

fish over dams, including elevators,

locks, and pool-type fishways. For

pool-type fishways, which consist of a

series of pools , each higher than the

next pool downstream, the difference

in pool elevation should be about 0.25

m. Pools should be at least 2.4 m

long, 2.4 m wide, and 1.2 m deep, but

the size should be increased if large

numbers of shad are expected to use

the fishway.

Although the present state of

engineering and biological knowledge

appears to be adequate for providing

upstream passage of adults, the problem of safe passage of young and

adults back downstream have not been

adequately solved. Fish must usually

pass through turbines or over spillways, causing considerable mortality.

Turbidity, Substrate, and Depth

Shad eggs exposed to suspended

sediment concentrations as high as

1,000 mg/l did not affect hatching

success (Auld and Schubel 1978). The

mortality of larvae was high at concentrations of suspended sediments

greater than 100 mg/l for 96 hours.

American shad may spawn at many

different depths and over a variety of

substrates. They seem to prefer areas

dominated by shallow water or broad

flats with a sand or gravel bottom

(Mansueti and Kolb 1953; Walburg 1960;

Leggett 1976). They also spawn in

deeper waters adjacent to shoals

(Stier, in press).

At sea, shad tend to remain near

the bottom during the day and move up

in the water column to feed at night

(Neves and Depres 1979). Adults have

been found at depths of 20-340 m, but

most occur at 50-100 m.

14
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PP*

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16
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Dep. 135 pp.

Ulrich, G.F., N.C. Jenkins, and J.W.

McCord. 1979b. Monitoring and

assessment of

shad fishery.

No. AFCS-7-1.

Resour. Dep.,

44 PP.

Ulrich, G.G ., N.C.

South Carolina's

Compl. Rep., Proj.

S.C. Wildl. Mar.

Mar. Resour. Div.

Jenkins, and J.W.

McCord. 1983. Monitoring and

assessment of the South Carolina

commercial fishery for American

shad. Compl. Rep., Proj. No.

AFCS-8-3. S.C. Wildl. Mar.

Resour. Dep., Mar. Resour. Div.

70 PP.

Vinogradov, V.I. 1981. Daily rhythms

and food rations of common

pelagic fishes in Nantucket

Shoals (New England) in the

summer. Soviet J. Mar. Sci.

(English translation of Biologiya

Morya) 7:170-174.

Walburg, C.H. 1956. Observations on

the food and growth of juvenile

American shad, Alosa sa idissima.

Trans. Am. Fish..&

Walburg, C.H. 1960. Abundance and

life history of the shad, St.

Johns River, Florida. U.S. Fish

Wildl. Serv. Fish. Bull. 60:487-

501.

Walburg, C.H., and P.R. Nichols.

1967. Biology and management of

the American shad and status of

the fisheries, Altantic coast of

the United States, 1960. U.S.

Fish Wildl. Serv. Spec. Sci. Rep.

Fish. No. 550. 105 pp.

Williams, R. O., and G. E. Bruger.

1972. Investigations on American

shad in the St. Johns River. Fla.

Dep. Nat. kesour. Tech. Ser. 66.

49 PP.

Williams, R., W. Gray, and J. Huff.

1975. Study of anadromous fishes

of Florida. Compl. Rep. NMFS

Grant-in-aid Program, Proj.

AFCS-5. Florida Dep. Nat.

Resour. Mar. Res. Lab. 160 pp.

1 8
p77 -101

REPORT DOCUMENTATION 1. M+uRT NO.

Biological Report 82(11.45)* I*.

3. -CW*W’S Acc,,,,~. ,,Q

PAGE

I. Title .M S,,bt,lh

Species Profiles: Life Histories and Environmental Requirements i R*eoon oat*

of Coastal Fishes and Invertebrates (South Atlantic)--American April 1986

L

Shad

L Author(*)

Douglas E. Facey and Michael J. Van Den Avyle

Georgia Cooperative Fishery and Wildlife Research Unit 10. l ~l.cff-f.~L/wOIc "nil "0.

School of Forest Resources

University of Georgia 11. *CnctfQ OI G,.n((C, NQ.

Athens, GA 30602 fcl

ml

12 %gOnUdrY O~mnl~tton Nama ati Addn,a

National Coastal Ecosystems Team U.S. Army Corps of Engineers :J. QPda*poRL~riod-"

Fish and Wildlife Service Waterways Experiment Station

U.S. Department of the Interior P.O. Box 631

Washington, DC 20240 Vicksburg, MS 39180 II.

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

Species profiles. are literature summaries of the life history, distribution, and

environmental requirements of coastal fishes and invertebrates. Profiles are prepared to

assist with environmental impact assessment. The American shad (Alosa sapidissima) was

the most valuable food fish on the Atlantic coast prior to the 19mbut landings have

declined drastically since that time, and-sport fishing has become more important than

commercial fishing in most rivers where stocks still exist. The American shad is

anadromous: adults congregate offshore in areas within the North Atlantic Bight during

summer, move southward in coastal waters during fall and early winter, and move up rivers

to spawn during late winter-hand spring. In rivers in the South Atlantic Region, the

spawning run peaks during January and February, but can continue through April and May.

Larval shad live in upriver areas and eat a variety of small invertebrates. Juveniles

gradually move downriver and live in estuaries, where they grow to about 100 mm in length

and migrate to sea where they live 3-5 years before maturing. Most American shad return

to their natal river to spawn;.and in the South Atlantic Bight, most adults die after

spawning once. Development of eggs requires water temperatures of 14 to 21 "C, dissolved

oxygen concentrations of at least 5 mg/l, and current velocities capable of buoying eggs

during incubation. Downstream transport of the young and spawning migrations of adults

can be adversely affected by excessive alteration of the river flow.

7. DoarWNmt AlWlyis a. O*scripton

Estuaries

Fishes

Growth

Feeding habits

. td9ntlfiWSf~n.C,,d.d T.,m‘

American shad

Alosa sapidissima

Habitat

Salinity requirements

Temperature requirements

Spawning

Fisheries

L ArmlabIlity ttatw”mt

Release unlimited

_ _____ ___ __ _ (tea ANSI-ZI4’J.lU~

U.S.G.P.O. 1986/661-638

7)

Species proiiies :

Facey, Dougias i.

*

x

l

\1L .

l

ncf n,~~h Library rd”s Department of the Inter-

‘>iitiinel Biological Sur”@J’

Headquarters D~ws,on of &&glCa,

SerViCeS. Washington, DC

Eastern Energy and Land “se Team

Leetown, WV

National Coastal ~~~~~~~~~~ Tag,,,

Slidall. LA

WeStern Energy and Land “se ~~~~

Ft Collins. CD

Locations of Regmnal Offices

REGION 1

Regional Director

U.S. Fish and Wildlife Service

Lloyd Five Hundred Building, Suite 1692

500 N.E. Multnomah Street

Portland, Oregon 97232

REGION 4

Regional Director

U.S. Fish and Wildlife Service

Richard B. Russell Building

75 Spring Street, S.W.

Atlanta, Georgia 30303

REGION 2 REGION 3

Regional Director Regional Director

U.S. Fish and Wildlife Service U.S. Fish and Wildlife Service

P.O. Box 1306 Federal Building, Fort Snelling

Albuquerque, New Mexico 87 IO3 Twin Cities, Minnesota 55 I I 1

REGION 5

Regional Director

U.S. Fish and Wildlife Service

One Gateway Center

Newton Corner, Massachusetts 02158

REGION 6

Regional Director

U.S. Fish and Wildlife Service

P.O. Box 25486

Denver Federal Center

Denver. Colorado 80225

REGION 7

Regional Director

U.S. Fish and Wildlife Service

1011 E. Tudor Road

Anchorage, Alaska 99503
DEPARTMENT OF THE INTERIOR

U.S. FISH AND WIlDLIFE SLRVICE

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 theenvironmental and cultural values of our national parks and historical places,

and providing for the enjoyment of life through outdoor recreetion. The Department assesses our energy and mineral resources end works to assure that their development is in

the best interests of all our people. The Department also has a major responsibility for

American Indian reservation communities and for people who live in island territories under

U.S. adminlstmtion.