Species Profiles: Life Histories and

Environmental Requirements of Coastal Fishes

and Invertebrates (Mid-Atlantic)

AMERICAN SHAD

QL

155

.S63

no. 82

11.37

L L

Fish and Wildlife Service

U.S. Department of the Interior U.S. Army Corps of Engineers

Coastal Ecology Group

Waterways Experiment Station

Biological Report 82(11~7)

April 1985

Library National Wetlands Reetaflh Center

11 c Ftsh and Wdllfe Sefllce

700 cajundome EWJ~

Lafayette, La. 70506 TR EL-824
This is one of the first reports to be published in the new "Biological

Report" series. This technical report series, published by the Research

and Development branch of the U.S. Fish and Wildlife Service, replaces

the "FWS/OBS" series'published from 1976 to September 1984. The Biological Report series is designed for the rapid publication of reports with

an application orientation, and it continues the focus of the FWS/OBS

series on resource management issues and fish and wildlife needs.
Biological Report 82(11.37)

TR EL-82-4

April 1985

Species Profiles: Life Histories and Environmental Requirements

of Coastal Fishes and Invertebrates (Mid-Atlantic)

AMERICAN SHAD

Chet MacKenzie

Lori S. Weiss-Glanz

and

John R. Moring

Maine Cooperative Fishery Research Unit

313 Murray Hall

University of Maine

Orono, ME 04469

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 coasts fishes and invertebrates. U.S. Fish

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

This profile should be cited as follows:

MacKenzie, C., L.S. Weiss-Glanz, and J.R. Moring. 1985. Species profiles: life

histories and environmental requirements of coastal fishes and invertebrates

(mid-Atlantic)--American shad. U.S. Fish Wildl. Serv. Biol. Rep. 82(11.37).

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 BY To Obtain

millimeters (mn) 0.03937 inches

centimeters (cm) 0.3937 inches

meters (m) 3.281 feet

kilometers (km) 0.6214 miles

square meters (m2) _

square kilometers (km‘)

hectares (ha)

10.76 square feet

0.3861 square miles

2.471 acres

liters (1) 0.2642

cubic meters (m3)

gallons

35.31 cubic feet

cubic meters 0.0008110 acre-feet

milligrams (mg) 0.00003527 ounces

grams (9) 0.03527 ounces

kilograms (kg) 2.205 pounds

metric tons (t) 2205.0 pounds

metric tons 1.102 short tons

kilocalories (kcal) 3.968 British thermal units

Celsius degrees 1.8("C) + 32 Fahrenheit degrees

inches 25.40 millimeters

inches 2.54 centimeters

feet (ft) 0.3048 meters

fathoms 1.829 meters

miles (mi) 1.609 kilometers

nautical miles (mii) 1.852 kilometers

square feet (ft')

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 (oz)

pounds (lb)

short tons (ton) . .

CONVERSION TABLE

Metric to U.S. Customary

U.S. Customary to Metric

28.35

0.4536

0.9072

British thermal units (Btu) 0.2520

Fahrenheit degrees 0.5556("F

iv

32) Celsius degrees

grams

kilograms

metric tons

kilocalories
CONTENTS

PREFACE ........................

CONVERSION TABLE ...................

ACKNOWLEDGMENTS. ...................

NOMENCLATURE/TAXONOMY/RANGE. .............

MORPHOLOGY/IDENTIFICATON AIDS. ............

REASON FOR INCLUSION IN SERIES ............

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

Spawning. ....................

Fecundity and Eggs. ...............

Larvae to Adults. ................

Ocean Migratiorl .................

GROWTH CHARACTERISTICS ................

THE FISHERY. .....................

History .....................

Current Statistics. ...............

Population Dynamics ...............

ECOLOGICAL ROLE. ...................

Foods ......................

Predators ....................

Diseases. ....................

ENVIRONMENTAL REQUIREMENT5 ..............

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

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

Oxygen ......................

Turbidity ....................

Substrate ....................

Depttt ......................

Water Movement. .................

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

V

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ACKNOWLEDGMENTS

The authors thank Christine M. Moffitt, Idaho Cooperative Fishery Research

Unit, University of Idaho; Richard J. Neves, Virginia Cooperative Fishery

Research Unit, Virginia Polytechnic Institute and State University; and Roy W.

Miller, Division of Fish and Wildlife, Dover, Delaware, for reviewing portions of

the manuscript.

vi
Figure 1. The American shad.

AMERICAN SHAD

NOMENCLATURE/TAXONOMY/RANGE

Scientific name . . Alosa sapidissima

(Wilson)

Preferred common name . . . American

shad (Figure 1).

Other common names . . shad, alose,

common shad, Atlantic shad, North

River shad, Potomac shad, Connecticut River shad, Delaware shad, Susquehanna shad, white shad, buck shad

(males only), poplarback shad (Scott

and Crossman 1973).

Class . . . . . . . . . Osteichthyes

Order . . . . . . . . . Clupeiformes

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

Geographic range: American shad are

anadromous. They are distributed

along the Atlantic coast from Newfoundland to Florida, and are most

abundant from Connecticut to North

Carolina. In the mid-Atlantic

region the American shad ascend

essentially all major rivers, but

abundance in some rivers is limited

by pollution or restricted by dams

(Figure 2).

On the Pacific coast, the

American shad was introduced into

the Sacramento and Columbia Rivers

in 1871, and the species is now

established from southern California

northward to Cook Inlet, Alaska, and

the Kamchatka Peninsula in Asia.

MORPHOLOGY/IDENTIFICATION AIDS

Body elongate, strongly compressed laterally, and rather deep,

its depth 17%-19% of total length (TL)

(Leim 1924; Bigelow and Schroeder

1953; Scott and Crossman 1973). Head

broadly triangular, 22%-24% of TL.

Gill membranes free from isthmus. Eye

moderate, adipose e.yelid well developed, diameter of eye 27%-32% of head

length (HL); snout moderate, length

27%-32% HL; interorbital width 19%-22%

of HL. The anterior end of the lower

jaw not especially thick or heavy and

somewhat pointed, and fitting easily

into a deep notch in upper jaw so that

the jaws are about equal when the

mouth is closed. The upper outline of

the lower jaw slightly concave. Maxillary extending to posterior margin of

eye. Teeth small, weak, and few in

1
NEW YORK

PHILADELPHIA

ATLANTIC OCEAN

3

e

N

I

MILES

OV

KILOMETERS

NORTH CAROLINA

CAPE HATTERAS

Figure 2. American shad are distributed along the coast of the Mid-Atlantic

Region from Massachusetts to North Carolina, but are most abundant in this

region along the coast from Connecticut to North Carolina. They ascend

essentially all major rivers but may be limited in some rivers by pollution

or restricted by dams.

2
number on premaxillary and mandible

(lost completely in the adult) and absent on the roof of the mouth. Gill

rakers on lower limb 59-73; branchiostegal rays 7,7 (rarely 7,6). Fins

soft rayed: dorsal-l, height moderate,

base short, ll%-13% of TL, rays 15-19,

usual1.y 17-15; caudal,

forked; anal-l,

distinctly

than dorsal

base length greater

base, 13%-14% of TL,

height shorter than dorsal

rays 18-24,

height,

usually 20-22; pelvics,

abdominal, small,

rays 9;

length 9%-10% of TL,

pectorals, low on sides,

length 14%-15% of TL, rays 14-18,

Iusually 16. Scales, large, crenulate

on posterior margin, deciduous.

Lateral line poorly developed with

about 50-55 scales. Ventral scutes

well developed forming a sharp sawbelly, prepelvic scutes 19-23, usually

20-22, postpelvic scutes 12-19, usually 15-17.

55-58.

Vertebrae 53-59, usually

Peritoneal lining pale; pyloric caeca numerous and usually c,lustered on right side. Usually 4 to 6

black spots in horizontal row behind

operculum. Size: average total length

380 mm; males up to 2.7 kg; females up

to 3.6 kg, rarely to 5.4 kg.

REASON FOR INCLUSION IN THE SERIES

Historically, commercial fishing

for American shad on the Atlantic

coast has been intensive and widespread. Now the fishery has virtually

collapsed and is important only in a

few rivers (Connecticut R.,

Connecticut; Hudson R., New York;

Neuse R., North Carolina; York R.,

Virginia) in the mid-Atlantic region.

In the rivers that support strong

runs, sport fishing for shad is more

important than commercial fishing.

Federal and State agencies have

initiated management programs aimed at

restoring American shad to their

former range and abundance. The

American shad is a natural resource

that should be considered for

protection in coastal and riverine

development projects.

LIFE HISTORY

Spawning

The American shad is an anadromous fish that lives several years in

the ocean and then returns to its

river of origin to spawn. At one

time, the species probably spawned in

virtually every accessible river and

tributary along the Atlantic coast of

North America.

The freshwater spawning migration

in winter, spring, or summer is timed

to correspond to favorable river water

temperatures. Shad usually migrate

far enough upstream so that the eggs

drift downstream and hatch before

reaching saltwater. In one study in

the Connecticut River, low river

temperatures may have delayed the

maturation of gonads causing the shad

to migrate farther upstream to spawn

(Marcy 1976). Glebe and Leggett

(1981) reported that the gonads of

American shad mature prior to entering

the Connecticut River.

American shad spawn as early as

mid-November in Florida and as late as

July in some Canadian rivers. Males

arrive at the spawning grounds before

females (Chittenden 1975). Water

temperature is the primary factor that

triggers spawning, but photoperiod,

flow velocity, and water turbidity

also exert some influence (Leggett and

Whitney 1972).

Although shad eggs in Virginia

streams become abundant after the

water temperature reach 12°C (Massman

1952), most spawning is from 13" to

20°C (Walburg and Nichols 1967). In

North Carolina, peak spawning is at

water temperatures near 20°C (Sholar

1976).

The diameter and abundance of

shad eggs collected in plankton nets

in the Pamunkey River, Virginia,

indicate that spawning takes place at

all hours of the night and day but is

3
more intense from noon to midnight

(Massman 1952). According to Miller

et al. (1971, 1975), spawning begins

in the evening between 1900 h and

2000 h and peaks between 2100 h and

2300 h in the Delaware River. After

0100 n, most of the spawning subsides.

On overcast days, spawning begins

earlier than on clear days, which

supports the hypothesis that the daily

onset of spawning is regulated by

light intensity (Miller et al. 1982).

Group spawning involving several

males and a female has been observed

(Marcy 1972). The spawning fish swim

vigorously at the surface, forming a

closely packed circle. Eggs are

released into the water and then

fertilized by the males (Marcy 1972;

Scott and Crossman 1973).

Shad spawn in main streams of

North Carolina rivers over sand shoals

where there is sufficient current to

keep the eggs suspended in the water

column (Sholar 1976). Shad spawn over

sand, silt, muck, gravel, and boulder

substrates (Mansueti and Kolh 1953;

Walh?rrg 1960; Leggett 1976). They

spawn in water depths about 1 to 10 m

(3 to 30 ft) but most often less than

3 m (10 ft) (Walburg and Nichols

1967). The current in spawning areas

ranges from about 0.5 to 3 ft/s

(Walburg and ,Nichols 1967; Marcy

1972).

The percentage of shad that spawn

more than once increases from south to

north (Table 1). For example, shad in

rivers south of Cape Fear, North

Carolina, die after spawning, but in

more northerly rivers some shad

survive spawning to spawn again. Shad

may return to spawn for up to 6

years (Table 2). The percentage of

repeat spawners ranges from 0% for the

St. Johns River, Florida, south of

Cape Fear, North Carolina, to 73% in

the St. John River, New Brunswick

(Carscadden and Leggett 1975a, 1975b).

The Delaware River, unlike other midAtlantic coast shad rivers, has few

repeat spawners (1.5% to 6.5%)

Table 1. The percent of repeat

spawners in American shad spawning

runs in 4tlantic coast rivers (Leggett

and Carscadden 1978).

---

River and Latitude Repeat

location of river ("N) spawners(%)

Miramichi (NB) 49 64

St. John (NB) 45 73

Connecticut (CT) 41 63

Hudson (NY) 41 57

Delawarea (DL) 40 6

Susquehanna (VA) 40 37

Potomac (VA) 38 20

York (VA) 37 24

James (V4) 37 27

Neuse (NC) 35 3

Edisto (SC) 33 0

Ogeechee (GA) 32 0

-

St. Johns (FL) -___~30 0

a Chittenden (1975).

according to Chittenden (1975). He

suggests that the probable causes for

the scarcity of repeat spawners there

are pollution and overfishing.

Fecundity and Ew

The American shad has a relatively high fecundity (116,000 to

659,000 eggs per female; Table 2).

Many eggs fail to fertilize and only a

small percentage of the fertilized

eggs hatch. High egg mortality has

been attributed to failure to fertilize, suffocation, fungus infection,

and predation (Leach 1925; Mansueti

and Kolb 1953).

The fecundity of spawners decreases from south to north (Table 3).

These trends in fecundity are independent of age and size.

Unfertilized eggs are irregularly

round in form and are about 1.8 mm in

diameter; fertilized, waterhardened

eggs are 2.5 to 3.5 mm in diameter and

4
Table 2. Lengths, weights, ages, and range of the number of eggs of American shad in the spawning populations of seven Atlantic coast rivers of the

United States, 1951-1959 (Walburg and Nichols 1967).

-~ -

Range of

Fork Body Age number of eggs

River length (mm) weight (kg) (years) per female

__--__

(x 1,000)

Hudson (NY) 355-556 0.8-3.0 3-9 116-468

Potomac (VA) 460-505 1.4-2.4 5-6 267-525

York (VA) 399-470 1.1-2.1 4-6 169-436

Neuse (NC) 447-498 1.8-2.7 4-6 423-547

Edisto (SC) 465-498 1.6-2.2 4-5 360-480

Ogeechee (GA) 457-475 1.7-2.2 4-6 359-501

St. Johns (FL)_ 368-460 0.6-1.8 4-6 277-659

- _ ____-_-____

Table 3. Mean virgin (first time

spawners) and lifetime fecundities

of American shad populations from

five Atlantic coast rivers, 1958-

1973 (Leggett and Carscadden 1978).

- -

Virgin Lifetime

River fecundity - -fecundity

St. Johns (FL) 406,000 406,000

York (VA) 259,000 327,000

Connecticut (CT) 256,000 384,000

St. John (NB) 135,000 273,000

Miramichi (NB) 129,000 258,000

are transparent, pale pink, or amber

(Marcy 1976; Miller et al. 1982).

Fertilized eggs are slightly heavier

than water and are nonadhesive. The

rate of development of shad eggs is

linearly related to temperature

(Mansueti and Kolb 1953). Eggs hatch

in 8 to 12 days at 11" to 15°C: 6 to 8

days at 17°C; and 3 days at 24°C

(Bigelow and Schroeder 1953; Scott and

Crossman 1973). No viable eggs

develop at water temperatures above

29°C (Bradford et al. 1966).

Larvae to Adults

American shad yolk-sac larvae are

about 6 to 10 mm long (TL) at hatching

and 9-12 mm TL when the egg yolk is

absorbed (Marcy 1976). Initially the

larvae 9-27 mm TL are planktonic

(Mansueti and Hardy 1967; Jones et al.

1978). They reach the juvenile stage

when about 25 to 28 mm long and about

4 weeks old (Jones et al. 1978).

Juveniles spend the first summer in

the river feeding on crustaceans and -

aquatic insects at the surface or in

the water column (Leim 1924: Walburg

1957; Levesque and Reed 1572).

In the fall, juveniles (75 to 125

mm long) migrate down the rivers to

brackish water and then to the sea. A

decrease in river water temperature

seems to trigger the migration

(Chittenden 1972). Juvenile shad

migrate seaward first in northern

rivers

southern

and progres;~v;~ttlater

rivers

Juvenile shad leave the St.

1977;:

Johns

River, Florida, as the water cools to

15.5OC (Walhurg 1960). In the

Delaware River, shad begin moving

downstream when the water temperature

drops to about 20°C; the movement

peaks at 15°C (Sykes and Lehman 1957).

In the Connecticut River, downstream

migration peaks late in September and

October. Peaks for the Upper Delaware

River and the Chesapeake Bay arc? late

October and late November, respectively. Once in the ocean, the shad

remain there until they mature.

5
Males become sexually mature when

they are 3 to 5 years old and the

females mature at 4 to 6 years old

(Leim 1924). Most of the male

first-time (virgin) spawners in the

Neuse (North Carolina) and Susquehanna

(Maryland) Rivers are 4 years old

(LaPointe 1958). The majority of

female virgin spawners are 4 to 5

years old. Shad are known to complete

their entire life cycle in only one

freshwater system: Millerton Lake, a

reservoir on the San Joaquin River,

California (Lambert et al. 1980).

Morphological characteristics for

distinguishing shad larvae from other

larvae of similar species are

described by Jones et al, (1978).

Ocean MigrationThe American shad in the midAtlantic region form large schools and

undertake extensive ocean migration

(Leggett and Whitney 1972). Shad from

all Atlantic coast rivers spend the

summer and fall in the Gulf of Maine

(Talbot and Sykes 1358). This

congregation includes immature shad

and spawned-out adults from rivers

north of Chesapeake Bay. Shad are

apparently scattered along the

mid-Atlantic coast during the winter

(Talbot and Sykes 1958; Walburg and

Nichols 1967). Migrating American

shad seek bottom water temperatures

between 3" and 15°C but probably

prefer temperatures between 7" and

13°C (Neves and Depres 1979).

In early spring, the schooyho;z

shad migrate toward the coast.

returning to rivers south of Cape

Hatteras follow the Gulf Stream t o

remain within the 3" to 15'C bottom

isotherm. Those migrating to rivers

north of Cape Hatteras later in the

spring follow a route farther seaward

into the Middle Atlantic Bight where

water temperatures have risen

sufficiently. Tag returns indicate

that some schools migrate north along

the coast (Neves and Depres 1979).

American shad migrate as much as

21 km/day in Chesapeake Bay andsot:;

Bay of Fundy (Leggett 1977).

migrate iup to 3,000 km during the

spring or fall migration.

A large majority of American shad

return to their natal river to spawn.

Homing behavior involves both olfaction and rheotaxis (Dodson and Leggett

1974). The homing mechanism is

sufficiently robust to perpetuate

migrations even after major changes in

water flow, such as below a

hydroelectric dam.

GROWTH CHARACTERISTICS

American shad live to be 5 to 7

years old and most weigh between 1 to

3 kg. The oldest shad reported for

the United States was 11 years of age

and 584 mm long (Scott and Crossman

1973). Factors that affect growth

sometimes can be identified by

examining fish scales. Slow growth in

the winter causes growth rings on

scales to be close together; these

winter marks (annuli) can be used to

age fish. Judy (1961) verified the

scale method for aging American shad.

He also noted a mark on the scales

that formed when the juveniles left

fresh water, and described marks that

formed at spawning. Shad grow about

100 mmlyr until sexually mature; after

maturity, growth slows (Table 4).

Table 4. The average total lengths of

shad and annual growth increments (mm)

for ages I to VII in the Bay of Fundy

(Leim 1924).

Growing

Season _ Age Length Increment - -

1st I 120

2nd II 240 120

3rd III 320 80

4th IV 400 80

5th V 470 70

6th VI 520 50

7th VII 570 50

6
THE FISHERY

-History

In the 19th century, extensive

fisheries for shad developed along the

entire Atlantic coast from the St.

Johns River, Florida, to the St.

Lawrence River, Canada. Major types

of gear were drift and staked gill

nets, pound nets, haul seines, weirs,

fyke nets, bow nets, and dip nets.

The estimated U.S. Atlantic coast

catch in 1896 was 22,680 metric tons

(50 million lb). Between 1930 and

1960, the average annual catch was

about 4,530 metric tons or 10 million

lb (Figure 3). In 1983, landings were

about 1,585 metric tons (3.5 million

lb).

Commercial shad landings in

Chesapeake Bay were traditionally the

largest along the Atlantic coast

(Table 5), but because of the closure

of Maryland's shad fishery in 1980,

the mid-Atlantic and South Atlantic

catches have been greater. Dams,

pollution, and overfishing have

contributed to the decline of shad

stocks.

Current Statistics

Rhode Island. American shad are

an incidental part of the commercial

catch in Rhode Island. Approximately

77,000 lb were landed in 1982 (Table

6). The Runnings and Warren Rivers

support a small sport fishery. Shad

AMERICAN SHAD - U.S. ATLANTIC COAST

25000 -

Figure 3. Landings of American shad, U.S. Atlantic coast, 1880-1976 (Walburg

and Nichols 1967;- updated and reproduced with oermission from Public Service

Electric and Gas Company, 80 Park Plaza, Newark, New Jersey 07101).

7
Table 5. The landings of American shad (x 1,000 lb) for different regions

along the U.S. coast, 1960-83. Code: NE=New England, MA=Mid-Atlantic, CB=

Chesapeake Bay, SA=South Atlantic, PC=Pacific Coast (1960-77, National Marine

Fisheries Service, Statistical Digest; 1978-83, National Marine Fisheries

Service, unpublished data).

-___ -

Year NE

Coastal regions -

MA CB SA PC

1960 432 1,237 2,682 1,614 456

1961 547 1,026 3,144 1,612 927

1962 470 841 3,795 2,167 1,586

1963 325 744 3,139 1,734 1,503

1964 320 721 3,541 1,687 818

1965 380 635 4,298 2,379 870

1966 279 379 3,564 1,736 1,347

1967 754 387 3,005 1,562 1,333

1968 218 379 3,508 2,052 862

1969 201 342 3,540 1,904 610

1970 186 314 5,151 1,851 724

1971 283 222 2,473 1,452 499

1972 264 375 3,014 1,091 709

1973 261 308 3,033 685 483

1974 257 294 1,789 655 511

1975 208 337 1,321 518 522

1976 412 322 1,006 320 481

1977 418 394 1,547 418 560

1978 361 245 1,322 976 545

1979 330 216 1,041 363 797

1980 253 406 998 839 277

1981 66 510 500 1,235 120

1982 403 757 590 1,033 429

1983 504 365 242 1,974 413

have been reintroduced unsuccessfully

in the Pawcatuck River, in an attempt

to restore a spawning population

there; the river is currently closed

to shad fishing.

Connecticut. The Connecticut

River supports a modest commercial

shad fishery. Most fishing is with

drift gill nets below Hartford at

night. Some fishing is done during

the day when the water is turbid.

Fyke, trap, and pound nets are not

allowed in the river during the shad

run. The primary market is for roe

(eggs); males (buck shad) have little

or no value. The State of Connecticut

carefully regulates commercial fishing

for shad on the Connecticut River.

The season is open to commercial

fishing from April 1 to June 15.

During the season, fishing is

prohibited from Friday sunset to

Sunday sunset. Monofilament gill nets

are prohibited and the gill nets used

must have a minimum stretch Imeasure of

5 inches. There are no size or sex

restrictions. License holders must

report their catch at the end of each

fishing season.

Sportfishing for shad in Connecticut is permitted from April 1 to

a closing date determined each year.
Table 6. Annual (1960-83) commercial landings (x 1,000 lb) of American shad

from Rhode Island, Connecticut, New York, New Jersey, Maryland, and North

Carolina (1960-77, National Marine Fisheries Service, Statistical Digest;

1978-83, National Marine Fisheries Service, unpublished preliminary data).

Year

1960

1961

1962

1963

1964

1965

1966

1967

1968

1969

1970

1971

1972

1973

1974

1975

1976

1977

1978

1979

1980

1981

1982

1983

Rhode

Island

3

4

7

2

3

4

23

5

2

6

12

42

14

2

7

6

3

1

1

1

2

1

77

23

Connecticut

421

463

456

301

278

352

242

240

212

190

173

241

249

258

247

165

392

392

332

306

207

325

283

424

New New

York Jersey

472 694

303 633

243 480

202 442

141 430

133 392

ai 242

113 248

126 241

136 188

106 195

73 141

103 263

157 143

164 122

196 122

186 100

1 194

2 160

a 148

114 292

58 259

73 334

33 112

Mary1 and

1,336

1,815

1,575

a27

890

1,343

1,133

867

958

1,292

1,039

953

957

597

220

184

110

77

a7

47

24

1

9

27

North

Carol ina

702

673

765

693

640

1,069

701

777

a42

719

953

680

468

321

369

241

167

120

402

278

199

352

412

380

Angling and scoop nets are permitted

in streams. The daily bag limit is

six fish.

New York. The Hudson River has

the only commercial American shad

fishery of note in New York. Gill

nets are the principal type of

commercial gear used. Staked gill

nets contribute about 70% of the gill

net catch and drift gill nets contribute 30%.

New Jersey. Most commercial

fishing (almost entirely with gill

nets) in New Jersey is concentrated in

Delaware Bay; there is no sport

fishery in the bay. The only sport

fishery begins about 90 mi up the

Delaware River near Trenton, New

Jersey. Shad from the Delaware River

have been stocked into the Raritan

9

River basin in an attempt to restore

the once abundant runs there:, however,

all attempts of restoration have

failed (Didun 1983).

Pennsylvania. There is no commercial fishery for American shad in

Pennsylvania, but major restoration

attempts are underway on the

Susquehanna and Schuylkill Rivers

where sport fishing for shad is

currently banned. The estimated

annual value of the sport fishery on

the Schuylkill River fishery will be

$1,200,000 if the goal of restoration

is achieved (Mulfinger and Kaufmann

1980). Estimates of the annual value

of the current sport fishery on the

upper Delaware River range from

$828,000 to $3,000,000 (Miller et al.

1982).
Delaware. The legal commercial

gear in Delaware is 5- to 6-inch

(stretch measure) gill nets. Fishing

is allowed from March until early May.

In the past 30 years, the fishery has

shifted from the lower Delaware River,

where drift gill nets were used, to

Delaware Ray, where anchored gill nets

predominate (Miller 1982). The

increase in the commercial catch of

American shad in the Delaware Estuary

in recent years gives some evidence

that the species is becoming more

abundant (Table 7). The increase is

attributed partly to recently

completed pollution abatement projects

in the lower Delaware River (Miller

1982). There is no sport fishery

except for Nanticoke River and Broad

Creek, tributaries to Chesapeake Bay.

The fishing season for the Delaware

waters of the Delaware Estuary is from

June 10 to February 1.

Maryland. A sharp decline in the

commercial catch of American shad in

Maryland began in 1973 (Table 6).

This led to the closing of all waters

in varyland to shad fishing with the

exception of the Potomac River (which

is regulated by the Potomac River

Fisheries Commission) and coastal

waters (Carter and 'rleinrich 1982).

Table 7. American shad catch (x 1,000

lb) and value (x 1,000 dollars) for

Delaware and Virginia (Miller 1982;

Jack Travelstead, Virginia Marine

Resources Commission; pers. comm.).

Date Delaware Virginia

Catch Value Catch Value - -

1960 38 6 1.349 234

1965 110 16 21955 307

1970 13 1 4,112 315

1975 19 4 1,137 309

1976 36 8 896 284

I977 75 15 1,469 498

1978 70 20 1,235 212

1979 95 15 994 235

1980

1;;

31 974 353

1981 87 499 141

1982 333 585 180

Virginia. The biggest American

shad fishery is along the Atlantic

coast of Virginia. In 1982, 585,000

pounds were landed with a value of

$180,028 (Table 7). Gill nets are the

primary gear used.

North Carolina. In North

Carolina, shad are caught by commercial and sport fishermen from late

January to the end of March. The

commercial fishery employs drift and

staked gill nets, pound nets, and

seines. The primary types of gear

used on the Neuse River, which has

the largest commercial catch, are

staked and drift gill nets (Hawkins

1980).

_~Population Dynamics

Recruitment into the shad fishery

is largely dependent on the size of

the spawning stock and environmental

factors that govern spawning success

and survival. About 85% of the

variation in the numbers of American

shad that spawn in the Hudson River in

any one year is largely dependent on

the number of spawners 5, 4, and 1

year earlier (Talbot 1954). About 64%

of the annual variation in the abundance of juvenile shad in the

Connecticut River from 1966 to 1973

was directly related to the number of

spawners; 22% was attributed to

environmental factors, principally

water temperature and river flow

during the run (Marcy 1976).

The rate of exploitation is a

major factor influencing changes in

the abundance of shad in the

Connecticut River (Leggett 1976).

Because of the high value of shad eggs

(roe), females in the spawning run are

the main targets of the fishery.

The number of adults that survive

the fishery in any one year is

directly correlated with the numbers

of fish produced in the next

generation. This relationship is

described by a stock-recruitment

equation:

1 0
q = N 0.7 (I - N/87)

e

where R is recruitment and N is the

parent stock in terms of egg numbers

(Leggett 1976). Average annual

mortality rates, calculated from

tag-recaptures data, were about 70%

for males and 71% for females from

1965 to 1973 in the Connecticut River

(Leggett 1976).

The age structure of American

shad returning to spawn in Delaware

and North Carolina consists primarily

of 4- and 5-year-old males and 5- and

6-year-old females (Table 8).

ECOLOGICAL ROLE

Foods

Apparently, young American shad

in rivers feed mostly in the water

b

column (Levesque and Reed 1972;

Domermuth and Reed 1980). Early shad

larvae feed mostly on cyclopoid

copepods and tendipedids (Levesque and

Reed 1972). The stomach contents of

Table 8. Age composition (%) of the

spawning population of American shad

Delaware

;Fhitttheenden

River, Delaware

1975), and the Neuse

River, North Carolina (Hawkins 1980).

Delaware North Carolina

Age Number % Age Number %

II

III

IV

V

VI

IV

V

VI

VII

Males

: 00 2'6

236 76:l

62 20.0

3 0.0

Females

225: 61.6 13.7

88 24.1

2 0.0

Males

III 16

IV 126 3;

V 151 46

VI 310 9

VII 3 1

Females

IV

V 444: 5:

VII 10 1

VIII 1 0

juvenile (35-85 mm FL x = 55 mm) shad

six Atlantic coast rivers (St.

A!hns, Florida; Ogechee, Georgia;

Neuse, North Carolina; Pamunkey,

Virginia; Hudson, New York;

Connecticut, Connecticut) suggested

that shad ate suitable organisms that

were most available (Walburg 1957).

In contrast, a study performed on the

Connecticut River, Connecticut, by

Domermuth and Reed (1980)

demonstrated that juvenile shad (Tl =

28 to 132 mm x = 63.3 mm) were

selective; i.e., most selected daphnia

(51%) and bosmids (20%), whereas

despite high abundance, copepods (8%)

and benthos (0.1%) were consumed in

small quantities.

After going to sea, juveniles and

adults feed on a variety of small

crustaceans, many of which are benthic

organisms. Copepods and mysids

constituted 90% of the diet of adult

shad in the Bay of Fundy (Leim 1924).

Adults also feed on small fishes,

euphausids, fish eggs, and amphipods

(Bigelow and Schroeder 1953; Scott and

Crossman 1973). Most shad have a die1

vertical migration that follows the

die1 migration of their principal

food, zooplankton (Neves and Depres

1979). In a study off the coast of

North Carolina, 'anchovies (Anchoa

hepsetus) were in 12 of the 15

juvenile shad examined (87-141 mm),

and 39 of 41 adults (Holland and

Yelverton 1973). The stomachs also

contained zooplankton under 5 mm

(Holland and Yelverton 1973). This

size was reported by Atkinson (1951)

as being too small for shad gill

rakers to retain.

Food was scarce in the stomachs

of shad migrating upstream to spawn

(Leim 1924). But in one experiment,

migrating prespawning shad placed in a

freshwater pond fed on artificial feed

(Atkinson 1951).

Adult shad fed on mayflies in

freshwater and their stomachs

contained the remains of small fish

(Chittenden 1976). Shad strike;

11
artificial lures when in freshwater, a

behavior unexplained.

Predators

Juvenile shad may be preyed upon

by a variety of predators in freshwaters (Scott and Crossman 1973), but

Leim (1924) was unable to demonstrate

predation by either American eels

(Anguilla rostrata) or striped bass

(Morone sax-in the Shubenacadie

River, Nova Scotia. Seals prey on

American shad, but adults probably

have few enemies, except for humans

(Scott and Crossman 1973).

Diseases

The American shad has the usual

complement of parasites and diseases.

Acanthocephala, parasitic copepods,

distomes, nematodes, and trematodes

have all been reported in or on shad

(Scott and Crossman 1973). In the

Connecticut River, the sea lamprey

(Petromyzon marinus) and freshwater

lampreys (Ichthyomyzon spp.) sometimes

attach to adult shad (Walburg and

Nichols 1967).

ENVIRONMENTAL REQUIREMENTS

Salinity

The American shad adapts readily

to either freshwater or seawater

during its anadromous migrations. The

adults may spend two to three days in

the estuary before entering the river

(Leggett 1976). One test was made in

this connection. Transfer of adult

shad from seawater to freshwater over

a 2.5 hour period caused physiologic

stress and a mortality of 54% (Leggett

and O'Boyle 1976).

Eggs are always deposited in

freshwater and are believed to be

intolerant of full-strength seawater.

Leim (1924) suggests that shad eggs

and larvae tolerate brackish water

with a salinity as high as 15 ppt.

Temperature

Spawning runs into rivers

(Columbia River, Washington;

Connecticut River, Connect.icutMassachusetts; St. Johns River,

Florida) at various latitudes on the

Pacific and Atlantic coasts of North

America peak at water temperatures of

15.5' to 2O.O"C (Leggett and Whitney

1972).

The rate of development of shad

linearly related

~~$erat~~e (Mansueti and Kolh ,953;:

Eggs hatch in 8 to 12 days at 11' to

15°C; 6 to 8 days at 17“C and 3 days

at 24°C. (Leim 1924; Bigelow and

Schroeder 1953; Scott and Crossman

1973). E!w stop developing when

water temperatures drop to 7°C (Leim

1924). Abnormalities develop when the

temperature rises to 22OC (Leim 1924),

and no viable larvae develop from eggs

at water temperatures above 29°C

(Bradford et al. 1966).

The American shad is intolerant

of cold water temperatures. The lower

thermal tolerance limit is about 2°C

but prolonged exposure to 4" to 6°C

may cause high mortality or stress

(Chittenden 1972). If young shad are

given a choice, they generally avoid

temperatures below 8°C and strongly

avoid temperatures below 5°C.

Chittenden concluded that cold water

releases below large impoundments may

curtail or destroy historical spawning

and nursery areas there.

Juvenile American shad attempt to

avoid excessively high or rapid

increases in water temperature. Tests

in tanks show that shad avoid

temperature increases of about 4°C

above the acclimation temperature

(Moss 1970). They did not attempt to

avoid changes of l"C, suggesting a

sensory threshold of between +l" and

+4"C above ambient temperature.

Field observations bear out the

laboratory findings of Chittenden

(1972) and Moss (1970). Migrations of

17
American shad in the ocean and

freshwater are closely tied to changes

in vfater temperature. Shad are most

frequently caught commercially in

ocean bottom temperatures of 7' to

13°C (Neves and Depres 1979).

Oxygen

The American shad require welloxygenated waters either in rivers or

in the sea. Dissolved oxygen levels

must be at least 4 to 5 mg/l in

headponds through which shad pass in

their migration (Jessop 1975). In the

laboratory, equilibrium is lost at

dissolved oxygen levels below 3 mg/l;

heavy mortality occurs at :evels below

2 mg/l; and all fish die at concentrations less than 0.6 mg/l

(Chittenden 1969). Shad eggs were

absent where the concentration of

dissolved oxygen was lower than 5 mg/l

(Marcy 1976). The oxygen LCso for

Connecticut River shad eggs is 2.0 to

2.5 mg/l (Carlson 1968).

Turbidity

Extensive dredging of the Hudson

River produced no measurable adverse

effects on shad abundance (Talbot

1954). Adult shad readily enter the

Shuebenacadie River in Nova Scotia,

where suspended sediment concentration

sometimes is 1 g/l (Leim 1924). In a

laboratory study, mortality of eggs

held in concentrations of suspended

sediments up to 1 g/l from fertilization to hatching did not differ

significantly from control groups

(Auld and Schubel 1978); however, the

survival of shad larvae exposed to

concentrations greater than 0.1 g/l

for 96 hours was sharply reduced.

Larvae apparently are much less

tolerant of suspended sediments than

eggs.

Substrate

Substrate type apparently is

unimportant to shad. They spawn in

the water column and the eggs are

carried downstream. American shad

have been observed to spawn successfully over silt, mud, sand, gravel,

and boulders (Plansueti and Kolb 1953;

Walburg 1960; Leggett 1976). Only

under the most adverse of conditions,

in which mud covered and smothered the

eggs, was substrate a problem.

Depth

American shad show little depth

preference in freshwater. Adults are

caught during spawning runs in all

parts of the river channel. Spawning

has been observed in rivers at depths

ranging from 0.45 to 7 m (Mansueti

and Kolb 1953; Walburg 1960; Kuzmeskus

1977).

Juveniles were found at depths of

0.9 to 4.9 m in the Connecticut River

(Marcy 1976). Abundance was related

to the distance upstream and not to

depth. During the day, 87% of the

juvenile shad caught in a gill net

:;req ;e",' the bottom at depths of 3.7

. . At night, all were caught

near the surface. At sea, shad are

near the bottom during the day and

disperse in the water column at night

(Neves and Depres 1979).

Water Hovement

Water velocity is critical to

shad because shad must negotiate river

currents and occasional fishways when

migrating upstream, and pass safely

over spillways while going downstream.

Adult shad migrating upstream are

reluctant to use traditional fishways,

probably because entrance widths,

depths, and flows are often unsuitable

(Walburg and Nichols 1967). Pool-andweir fishways, vertical-baffle

fishways, and elevators are better

carriers of American shad. For the

pool-and-weir fishway, optimum difference in pool elevations is 23 cm when

water velocities are 61 to 91 cm/set.

For any fish passage to work, proper

current at the entrance is essential.

In the Connecticut River, the

daily movement upstream is about 5 km

.3
in brackish water, and 14 km in freshwater (Leggett 1976). Adult

downstream migration depends on water

currents and the pattern of currents

around obstructions. In the

Farmington River, a tributary of the

Connecticut River, flow rate into the

fishway accounts for 60% of the

variation in the number of downstream

migrants entering the fishway (Moffitt

1979). No fish entered the tributary

because of the lack of an adequate

attractant flow. Fish that fail to

find the downstream passage must pass

over the spillway or through

hydroelectric turbines. Between 57%

and 80% of juvenile shad that pass

through a 850 kw Ossberger turbine are

killed outright, and others may die

later of stress or are easy prey to

predators (Gloss 1982). A fishway may

cause mortality because of excessive

loss of scales and injury. At least

25% of the shad die as a result of the

fish-lift at Mactaquac, New Brunswick

(Jessop 1975). Exposure to lethal

nitrogen supersaturated waters below

the dam sometimes causes stress or

mortality (MacDonald and Hyatt 1973).

Spawning takes place in water

velocities ranging from 9.5 to 132

cm/set based on hydrographic data at

sites where Kuzmeskus (1977) found

fresh spawn. Spawning normally takes

place at velocities of 30 to 90 cm/set

(Walburg 1960).

14
LITERATURE CITED

Atkinson, C.E. 1951. Feeding habits

of shad (Alosa sa idissima) in

fresh water. --4__T Ecology 32 3):556-

557.

Auld, A.H., and J.R. Schubel. 1978.

Effects of suspended sediment on

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Coastal Mar. Sci. 6(2):153-164.

Bigelow, H.B., and W.C. Schroeder.

1953. Fishes of the Gulf of

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Fish. Bull. 53. 577 pp.

b Bradford, A., J. Miller, and K. Buss.

1966. Bio-assays on eggs and

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(Alosa sapihissima). Pages 52-60

in. Carlson.

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Suitability of

Susquehanna River for

restoration of shad. U.S. Dep.

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Wildl. Washington, @.C.

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shad, Alosa sapidissima: evidence

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Fish. Res. Board Can. 32(5):653-

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35 PP.

15
Dodson, J.J., and W.C. Leggett. 1974.

Role of olfaction and vision in

the behavior of American shad

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Connecticut River from Long Island Sound. J. Fish. Res. Board

Can. 31(10):1607-1619.

Domermuth, R.B., and R.J. Reed. 1980.

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migration. Can. J. Fish. Aquat.

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low-head hydropower sites. Page

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sa idissima). J. Fish. Res. Board Ci!hTpp422-1426.

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Friedersdorff, and R.W,

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Div. Fish Wildl. 13 pp.

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of American shad, Alosa sapidis17
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fish-ladder, Farmington River,

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temperature changes. Trans. Am.

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Presented to The Academy o f

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September 24 and 25, 1980.

National Marine Fisheries Service.

1960-1977. Statistical Digest,

Fishery Statistics of the United

States. 1J.S. Dep. Commerce.

Washington, DC.

Neves, R.J., and L. Depres. 1979. The

oceanic migration of American

shad, Alosa sapidissima, along

the At-i-%?% coast. U.S. Natl.

Mar. Fish. Serv. Fish. Bull.

77(1):199-212.

Scott, W.B., and E.J. Crossman. 1973.

Freshwater fishes of Canada.

Fish. Res. Board Can. Bull. 184.

966 pp.

Sholar, T.M. 1976. Status of American

shad in North Carolina. Pages

17-31 in Proceedings of a work -

shop on American shad, Dec.

14-16, Amherst, MA. U.S. Fish

and Wildl. Serv. and Natl. Mar.

Fish. Serv.

Stier, D.J. Habitat suitablity index

models: American shad. U.S. Fish

Wildl. Serv. In press.

Sykes, J.E., and B.A. Lehman. 1957.

Past and present Delaware River

shad fishery and considerations

for its future. U.S. Fish Wildl.

Serv. Res. Rep. No. 46. 25 pp.

Talbot, G.B. 1954. Factors associated

with fluctuations in abundance of

Hudson River shad. U.S. Fish

Wildl. Serv. Fish. Bull. 56:373-

413.

Talbot, G.B., and J.E. Sykes. 1958.

Atlantic coastal migrations of

American shad. U.S. Fish Wildl.

Serv. Fish. Bull. 58(142):473-

490.

Walburg, C.H. 1957. Observations on

the food and growth of juvenile

American shad,.Alosa sapidissima.

Trans. Am. Fish. sot. 86:

302-306.

Walburg, C.H. 1960. Abundance and

life history of the shad, St.

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Wildl. Serv. Fish. Bull. 60(177):

487-501.

Walburg, C.H., and P.R. Nichols. 1967.

Biology and management of the

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Fish. 5!50:1-105.

18
b

REPORT DOCUMENTATION ;I. REPORT ~0.

PAGE i Biol. Rep. 82(11.37)* i *.,

I I

4. nt1* and Subtitl. 5 R*Llorl oat*

Species Profiles: Life Histories and Environmental Require- April 1985

ments of Coastal Fishes and Invertebrates (Mid-Atlantic)-- a

Bmpriran shad

r. Author(~)

cc

a Pe+anninc OrSanization Rept. h

1. PWlOrminS OrSanlraWn Name and Address 10. ~mi9CI/Tash/Worh Unit No.

Maine Cooperative Fishery Research Unit University of Maine

U.S. Fish and Wildlife Service Orono, ME 04469 11. bnlr.Ct(C) or Grant(G) No.

313 Murray Hall (C)

It tponrorlry O~~nix~tlon N8meand Addms -(G)

13. TYtm d R9p.M 6 Period coven

National Coastal Ecosystems Team U.S. Army Corps of Engineers

Fish and Wildlife Service Waterways Experiment Station

U.S. Dep. of the Interior P.O. Box 631 14.

Washington, DC 20240 . Vicksburg, MS 39180

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

IL Abatnti Kimit: 200 words)

Species profiles are literature summaries on the taxonomy, morphology, distribution, life

history, and environmental requirements of coastal aquatic species. They are designed to

assist in environmental impact assessment. The depleted populations of the American

shad, Alosa sapidissima, are being restored in many of the rivers along the east coast

that originally supported large runs. The American shad is an anadromous fish that lives

several years in the ocean and returns to its natal river to spawn in the spring when

temperatures reach 12 "C. The eggs are carried by ctirrents downstream from spawning

sites in large rivers for 8-12 days until they hatch. The larvae, which metamorphose to

juveniles in 3-4 weeks, remain in the river until fall when they migrate to the sea.

Shad move offshore and southward during winter at water temperatures of 3-15 "C.

American shad feed on zooplankton. They adapt readily to fresh or saltwater, but they

prefer salinities exceeding 4 ppt.

Estuaries

Shad

Growth

Feeding

b. IdW.tlRen/O‘w,-Ended T.rmr

American shad Salinity requirements

Alosa sapidissima Temperature requirements

Habitat Spawning

Fisheries

C. COSATI Ficld/Gmup

i Ar8ilabllity Statwncnt

Unclassified

(Se. ANSl-239.lSl OPTIONAL FORM 272 M-77)

(Formerly NTl%35)

oep.rtment oi comm*re=

Species proiiies :

kacicenz ie, Chet ,

s,esc wcn mvrury

LJ. S. Department of the Interi’

Yational Biological Survey

outhern Science Center

Eastern Energy and Land Use Team

Leetown. WV

National Coastal Ecosystems Team ^. ~~I. .

w__ J 7 “-p--_c l---i

Western Energy and Land “se Team

Ft Collins. CO

Locations Of RegIonal 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, SW.

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 87 103 Twin Cities, Minnesota 55 I I I

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

101 I E. Tudor Road

Anchorage, Alaska 99503
DEPARTMENT OF THE INTERIOR

U.S. FISH AND WILDLIFE SERVICE

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 recreation. The Department assesses our energy and mineral resources and works to assure that their development is in

the best interests of ail 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. administration.