REFERENCE COPY 

Do Not Remove from the Library I* P r.1 t \ . Il. C C 

. . Biological Report 82(11.38) Na~io~~, , an IY d*" s Research ' v'Ck Center TR EL-82-4 

Aprll 1888 

700 Cajun Dome Boulevard 

Lafayette, Louisiana 70506 

Species Profiles: Life Histories and 

Environmental Requirements of Coastal Fishes 

and Invertebrates (North Atlantic) 

ATLANTIC HERRING 

Coastal Ecology Group 

Fish and Wildlife Service Waterways Experiment Stat ion 

U.S. Department of the Interior U.S. Army Corps of ~nginees 
Biological Report 82(11.38) 

TR EL-82-4 

April 1986 

Species Prof i 1 es: Life Histories and Environmental Requirements 

of Coastal Fishes and Invertebrates (North Atlantic) 

ATLANTIC HERRING 

Kevin H. Kelly 

and 

John R. Moring 

Maine Cooperative Fish and Wildlife 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 

Divi si on of Bi 01 ogi cal 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 requi rements of coasts 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: 

Kelly, K.H., and J.R. Moring. 1986. Species profiles: life histories and 

environmental requi rements of coastal fishes and invertebrates (North 

Atlantic)--Atlantic herring. U. S. Fish Wildl . Serv. Biol . Rep. 82(11.38 ). 

U.S. Army Corps of Engineers, TR EL-82-4. 22 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, 1 i fe hi story, ecological role, environmental 

requirements , and economic importance, if appl icabl e. A three-ri ng 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 Special i st 

National Coastal Ecosystems Team 

U.S. Fish and Wildlife Service 

NASA-Sl idel 1 Computer Compl ex 

1010 Gause Boul evard 

Slidell, LA 70458 

U.S. Army Engineer Waterways Experiment Station 

Attention: WESER-C 

Post Office Box 631 

Vicksburg, MS 39180 
CONVERSION TABLE 

Metric to U.S. Custmary 

Mu1 tiply & To Obtain 

meters (m) 

ki 1 meters ( bn) 

0.03937 inches 

0.3937 inches 

3.281 feet 

0.62 14 mil es 

2 

square meters (m ) 10.76 

square ki 1 meters ( bn2) 0.3861 

hectares (ha) 2.471 

liters (1) 

cubic meters (m3) 

cubic meters 

square feet 

square miles 

acres 

gal 1 ons 

cubic feet 

acre- feet 

mil 1 igrams [mg) 0.00003527 ounces 

grams (g) 0.03527 ounces 

kilograms (k ) 2.205 pounds metrlc tons ft) 2205.0 pounds 

metric tons 1.102 short tons 

ki 1 ocal ori es ( kcal ) 3.968 British thermal units 

Celsius degrees 1.8("C) + 32 Fahrenheit degrees 

U.S. Customary to Metric 

inches 25.40 

inches 2.54 

feet (f t) 0.3048 

fathoms 1.829 

miles (mi) 1.609 

nautical miles (nni) 1.852 

square feet (ft2) 

acres 2 

square miles (mi ) 

gal 1 ons (gal ) 

cubic ,feet (ft3) 

acre-feet 

ounces (02) 28.35 

pounds (lb) 0.4536 

short tons (ton) 0.9072 

British thermal units (Btu) 0.2520 

mil 1 imeters 

centimeters 

meters 

meters 

kil meters 

ki 1 meters 

square meters 

hectares 

square kil meters 

1 i ters 

cubic meters 

cubic meters 

grams 

ki 1 ograms 

metric tons 

ki 1 ocal ories 

Fahrenheit degrees 0.5556("F - 32) Celsius degrees 
CONTENTS 

PREFACE ................................. CONVERSIONTABLE ............................ ACKNOWLEDGMENTS ............................. 

NOMENCLATURE/TAXONOMY/RANGE . . MORPHOLOGY/IDENTIFICATION AIDS 

Distinctive Characteristics . REASON FOR INCLUSION IN SERIES 

LIFE HISTORY ......... Spawni ng .......... Fecundity and Eggs ..... Larvae ........... Juveniles .......... Adults ........... Growth Characteristics ... THE FISHERY .......... History of the Fishery ... ECOLOGICAL ROLE ........ ENVIRONMENTAL REQUIREMENTS . . Temperature ......... Sal i ni ty .......... Oxygen ........... Sediments and Turbidity ... Water Movement ....... Environmental Contaminants . 

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17 
ACKNOWLEDGMENTS 

We thank David Stevenson, Maine Department of Marine Resources, West 

Boothbay Harbor, Maine, and Vaughn C. Anthony, National Marine Fisheries Service, 

Woods Hole, Massachusetts, for reviewing the manuscript. 
Figure 1. At1 antic herri ng (Cl upea harengus L. ) . 

ATLANTIC HERRING 

Scientific name . . . Clupea harengus 

(Linnaeus) (American Fisheries 

Society 1980); Western North 

Atlantic populations of this fish 

are recognized as a subspecies in 

most current American and Russian 

1 i terature , but as a di sti nct 

species (Cl upea harengus) in Europe. 

The taxonomy of Clupeidae was 

extensively reviewed by Svetividov 

(1952). 

,Preferred common name . . . Atlantic 

herring (Figure 1) 

Other common names . . . Sea herring, 

Labrador herring , sardine , sperl i ng , bri t (Bigel ow and Schroeder 1953), 

herring (Hi ldebrand 1963). 

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

Order . . . . . . . . . . Cl upei formes 

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

Geographic range: Atlantic herring in 

Northwest Atlantic waters range from 

Greenland to North Carolina. They 

are most abundant north of Cape Cod, 

and relatively scarce south of New 

Jersey. In Europe, where certain 

races enter brackish water, herring 

range from Spi tsbergen and northern 

Norway south to the Strait of 

Gibraltar (Hildebrand 1963). 

MORPHOLOGY/IDENTIFICATION AIDS 

The following information is from 

Hildebrand (1963) and Mansueti and - 

~ardy - (1967). Fin rays: dorsal 

16-20, usually 17-19: anal 16-20, 

usually 17 or 18; pectoral 17-19. 

Scales between gi 11 openings and 

caudal base ca. 56-62; ventral scutes 

weakly developed, ca. 39-46; vertebrae 

55-57 (49-60: Mansueti and Hardy 

1967), gill rakers on lower limb of 

first arch 37-52. Proportions as 

percent of standard length (SL) (may 

include some juveni les): body depth 

20.0%-25.8%, head 1 ength 22.6%-26.4%; 

eye diameter 5.3%-7.7%. Body 

compressed. Maxi 1 lary rounded 

posteriorly, not quite reaching middle 

of eye, its margin with minute serrae; 

teeth on lower jaw, tongue, and in 

oval patch on vomer. Dorsal origin 

about midway along trunk, usually 

closer to caudal base than to tip of 

snout. Body iridescent, bluish above, 

silvery on sides and belly. Maximum 

total length (TL) about 450 mm, but 

most average about 300 mm. 
Distinctive Characteristics 

The fol lowing conspicuous field 

marks (Hi 1 debrand 1963) distinguish 

postlarval and adult Atlantic herring 

from alewives (A1 osa pseudoharengus) 

and shads (Alosa spp. ): the point of 

origin of the dorsal fin is about 

midway a1 ong the trunk (considerab-ly 

farther back than in Alosa), the body 

is not as deep, and the sharp mid1 ine 

of the belly is less strongly serrate. 

The Maine fishery has thrived for 

over a century and is still a valuable 

resource. For each $1 worth of 

herring landed in Maine, an income of 

$1.47 is generated within the State's 

economy (Briggs et al. 1982). The 

average annual value of yaine herring 

landings was $4.3 million in 1976-1982 

(Table 1). Herring are also important 

prey of such comercial species as 

Atlantic cod, pol lock, haddock, silver 

hake, red hake, white hake, swordfish, 

and bl uefin tuna. 

REASON FOR INCLUSION IN SERIES 

LIFE HISTORY 

Atlantic herring are commercially 

important in the western North 

Atlantic, particularly in the Gulf of 

Maine and adjacent Nova Scotian she1 f 

waters (Fogarty and Clark 1983). The 

main fishery for adults is 

concentrated in the Jeffreys Ledge 

(Figure 2) area of the western Gulf of 

Maine. The Georges Bank fishery, 

which peaked in 1968 at 374,000 t, 

collapsed in 1977 because of heavy 

fishing intensity from foreign .nations 

and has shown no signs of recovery 

since. The main fishery for juvenile 

herring, or "sardines," is in the 

coastal waters of Maine and New 

Brunswick. 

Spawn i ng 

Time of spawni n . Atlantic herring spawn in the Gu f of Maine once a 

year, usually in late summer or fall 

(late August to early November). The 

spawning period is protracted in some 

years (Anthony and Waring 1980b). 

Bigelow and Schroeder (1953) reported 

that the heaviest spawning in the 

Grand Manan Island and Machias Bay 

areas took place in July, August, and 

September. Spawning in coastal waters 

to the south and west occurred 

progressively later and was briefer in 

duration: from mid-August to October 

Table 1. Weights (thousands of pounds) and values (thousands of dollars) of 

herring landings by State, 1976-82. Data for 1976 are from National Marine 

Fisheries Service (1980) ; data for 1977-82 are prel iminary and unpubl ished. 

New Massa- Rhode ConnectMaine chusetts Island icut 

Year Wt. Val. iP%F wt. Val. Wt. Val. Wt. Val. 

"NA indicates data were not available. 
Figure 2. Gulf of Maine showing Atlantic herring spawning grounds as discussed 

in text. 
around Petit Manan Island and near 

Mount Desert Island, Maine; in October 

in Ipswich and Massachusetts Bays, 

Massachusetts; and in late October and 

early November near Woods Hole, 

Massachusetts. 

In 1960-65, spawning on Georges 

Bank and off Nova Scotia began in 

August and peaked in October. In the 

coastal Gulf of Maine, spawning began 

in August or September and peaked in 

either September or October, depending 

on the year (Boyar 1968). 

Recent 1 arval herring surveys 

indicate that herring begin spawning 

in late August and early September in 

waters of eastern Maine and in mid- to 

late September in midcoastal and western Maine (Graham 1982). Co I 1 ecti ons 

of yo1 k-sac 1 arvae indicated that 

spawning on Georges Bank-Nantucket 

Shoal s took place from mid-September 

to late December (Lough et al. 1979). 

Spawning of Atlantic herring has 

been reported in spring in the Gulf of 

Maine, but it is apparently 

unsuccessful (Bigelow and Schroeder 

1953; Boyar 1968; Si ndermann 1979; 

Anthony and Waring 1980b). Spring 

spawning has not been reported for 

Georges Bank (Boyar 1968; Si ndermann 

1979). 

Spawning sites. Several offshore 

spawning sites were found near the 

perimeter of Georges Bank (Boyar et 

a1 . 1973). Spawning concentrations of 

herring on Georges Bank in 1962 were 

reported to be as 1 ong as 64 to 80 km 

and wide as 6 to 13 km (Yudanov 1966). 

In the Gulf of Maine, spawning 

concentrations are less dense and the 

number of spawning sites has decreased 

in recent years. Recent larval 

surveys have indicated that the 

important spawning sites are located 

in Machias Bay along the eastern coast 

of Maine and in waters south of 

Boothbay Harbor and off Saco Bay 

(Graham 1982). As judged by the 

numbers of spawning and near-spawni ng 

adults, Jeffreys Ledge appears to be 

the most important spawning ground 

(Boyar et al. 1973). Other Atlantic. 

herring spawning sites in the Gulf of 

Maine area are Lurcher Shoals and 

Trinity Ledge off Nova Scotia, and 

Grand Manan off New Brunswick, 

Matinicus Rock and Pumpkin Ledge off 

Maine, and Stellwagen Bank off 

Massachusetts (Sindermann 1979; Figure 3\ 

Spawning habits and depth. Atlantic herrina are coastal oceanic 

spawners. bing the act of spawning, 

the females press close to the bottom 

and deposit ribbons of adhesive eggs 

on the substrate (Blaxter and Holl iday 

1963). The males swim above the 

females and release milt into the 

water. 

Spawning depths range from 10 to 

100 m in the Gulf of Maine (Sindermann 

1979), although herring have been 

reported to spawn as deep as 200 to 

240 m in European waters (Blaxter and 

Holl iday 1963). Herring in the Gulf 

of Maine and the Bay of Fundy were 

reported to spawn at depths of about 4 

to 110 m (Bigelow and Schroeder 1953), 

both inshore along the coast and on 

shoals and ledges as far as 8 to 40 km 

offshore. Depths of two spawning 

sites on Jeffreys Ledge (western Gulf 

of Maine) in 1974 were 30 to 45 m and 

40 to 50 m (Cooper et a1 . 1975). 

Egg beds in Georges Bank in 1970 

were in water 50 m deep (Caddy and 

Iles 1973). A spawning bed observed 

by Ti bbo et a1 . (1963) in the Gul f of 

St. Lawrence, Canada, was only 2 to 

6 m deep and eggs in Miramichi Bay, 

New Brunswick, were 0.9 to 4.3 m deep 

(Pottl e et a1 . 1980). Herring spawned 

from close inshore to depths of 9.1 to 

11.0 m in Newfoundland waters (Tibbo 

1956). In areas where onshore winds 

caused turbulent surface water near 

shore, herring spawned i n deeper 

waters. 

Spawning substrate. A1 though 

At1 antic herri na usual 1 v sDawn over 

gravel or rock, -vegetation and shells 
may also be used (Sindermann 1979). 

Underwater observations on Jeffreys 

Ledge revealed that 80% to 90% of the 

eggs adhered to the red alga Ptilota 

serrata, and the rest stuck to the 

upper surfaces of rocks and boulders 

(Cooper et al. 1975). Two types of 

spawning substrate were boul der/rock 

substrate with vegetation and coarse 

fragments of sand, gravel , and shel 1s 

without vegetation. Three of the six 

spawning areas observed were composed 

of rough boulder/rock substrate; slope 

gradients ranged from O0 to 40°. 

In Georges Bank waters, herring 

eggs were found only over a flat 

surface of fairly well-sorted, rounded 

glacial gravel, 2 to 10 mm in 

diameter; no eggs were seen on sand or 

shel 1 fragments (Caddy and Iles 1973). 

In Miramichi Ray, New Brunswuck, scuba 

divers reported that eggs were 

deposited on a1 gae--primari ly-~hondrus 

crispus, Palmeria palmata, Phyl lophora 

sp. , and Fucus sp. --with an apparent 

 reference for the last two of these 

taxa (Pottle et al. 1980). In waters 

near Newfound1 and, herring usual ly 

spawn on gravel or rock substrates 

where vegetation is abundant, though a 

few spawn on bare rock along sandy 

shores (Ti bbo 1956). 

Spawni ng temperatures and sal in- 

- i ti es. Duri ng spawning , water 

temperatures on two Jeffreys Ledge 

spawning beds ranged from 9 to 12 C 

(Cooper et al. 1975). In the eastern 

Gulf of Maine and Grand Manan Island 

area, spawning temperatures of 8 to 

11 OC were reported. Autumn spawners 

in European waters prefer temperatures 

of 8 to 15 OC (Blaxter and Holliday 

1963). 

Salinities in waters where 

herring spawn in the Gulf of Maine are 

normally near 33 ppt; Atlantic herring 

never spawn in brackish waters of the 

gulf (Bigel ow and Schroeder 1953). 

Water currents. A water current 

of at least 1 to 2 km/h is usually 

1979). Bottom water currents 

throughout a tidal cycle on the 

Jeff reys Ledge spawni ng grounds ranged 

from 0 to 2 km/h and averaged 0.3 to 

0.5 km/h (Cooper et al. 1975). 

Fecundity and Eggs 

Estimates of the average number 

of eggs per female for Gulf of Maine 

herring indicate an increase in 

fecundity in recent years (Table 2). 

This increase may be compensation for 

a decrease in stock size (Anthony and 

Waring 1980b). For Gulf of Maine 

herri ng , fecundity is closely re1 ated 

to the weight of the fish, derived by 

the rqation of the curvilinear form 

F = aW , where a = 40.39 and b = 1.413 

(Morse and Morris 1981). The relation 

between fecundity and total length of 

the fish is debermined by using the 

forqula F = aL where a = 1.714 x 

10- and b = 4.j56 (Morse and Morris 

1981). 

A relatively constant number of 

eggs seem to be produced by fish of a 

particular length but some eggs may be 

reabsorbed during maturation. In 

Table 2. Estimates of fecundity 

(thousands of eggs) for Atlantic herring of different lengths in the Gulf 

Year 

of Mai ne. 

Total 

length (cm) 1963-64a 1980b 1982' 

a~erki ns and Anthony (1969). 

b~orse and Morris (1981). C Kelly (1983). present during spawni ng (Si ndermann 
European herring, annual variations in 

fecundity have been related to 

differences in food intake (Hempel 

1971; Bowers and Holl iday 1960). Low 

fat content in females, caused by a 

scarcity of food, can reduce the 

number of eggs produced (Hempel 1971). 

Absol ute fecundity probably increases 

with a decrease in stock density. For 

example, fecundity of the Georges Bank 

herring each year was inversely 

related to population abundance in the 

previous year (Anthony and Waring 

1980b). 

Egg diameter after ferti 1 ization 

is typically 1.0 to 1.4 mm (Bigelow 

and Schroeder 1953; Mansueti and Hardy 

1967). Egg size at maturity may vary 

in different areas and at different 

times of the year (Bowers and Holliday 

1960; Hempel 1971). In the southern 

Gulf of St. Lawrence, mean egg weights 

were 12.2 mg in spring and 7.5 mg in 

fall; fecundity of fall spawners was 

about 50% higher than in spring 

spawners (Messi eh 1976). 

Egg incubation periods f 1 uctuate 

with temperature. Periods reported in 

the Gulf of Maine are 10 to 15 days 

(Bi gel ow and Schroeder 1953). 

Hatching occurs after 8-9 days at 

10 OC (Cooper et a1 . 1975). 

Size of egg beds. On Jeffreys 

Ledge in 1974, the area of egg beds 

ranaed from 700 to 1.300 me. and each 

bed- encompassed a distinct .and we1 1 - 

defined area (Cooper et al. 1975). 

Off Nova Scotia, a spawning area 

occupying 67,500 m2 was reported by 

McKenzie (1964), but he did not 

indicate whether the bed was 

continuous. A spawning area of 

375,000 mZ off the Canadian coast was 

reported by Tibbo et al. (1963). 

Thickness of layers of eggs. The 

number of egg layers in herring egg 

beds varies with the intensity of 

spawning. One egg bed on Jeffreys 

Ledge was composed of 3 to 15 layers 

and another contained 1 to 4 layers 

(Cooper et a1 . 1975). Egg 1 ayers were 

1 to 2 cm thick on Georges Bank (Caddy 

and Iles 1973) and around 3.25 cm 

thick on Trinity Ledge off southwest 

Nova Scoti a (McKenzi e 1964). Baxter 

(1971) reported that the presence of 

three to four layers is not harmful to 

the survival of eggs in the lower 

1 ayers ; however, rates of development 

are not uniform throughout the egg 

layers, and egg survival is lowest in 

the bottom layers (Baxter 1971). 

According to Hempel and Schubert 

(1969), if the eggs mass is thicker 

than three or four layers, high 

mortality occurs in the deeper layers. 

Larvae 

Atlantic herring larvae are 5 to 

9 mm long at hatching (Blaxter and 

Holliday 1963). The newly-hatched 

1 arvae become concentrated in the 

upper water column but are soon 

dispersed by surface and near-surface 

water currents (Lough 1975). An 

exception is yolk-sac larvae at 

spawning sites with vegetation; the 

larvae may stay in the vegetation for 

several days (Cooper et a1 . 1975). An 

estimated 75% of Atlantic herring 

larvae die within 4 days after 

hatching (Graham and Chenoweth 1973). 

Larval food. The larvae depend 

on the yo1 k sac for food during their 

first few days of life, but the yolk 

sac is absorbed when the larvae are 8 

to 12 mm SL (Blaxter and Holl iday 

1963). Some larvae begin feeding even 

before the yolk is fully absorbed. 

Larvae reared at 10 OC began feeding 2 

to 3 days after hatching (Lough et al. 

1982). Newly hatched larvae (7 to 

20 mm long) prey principal ly on 

copepod naupl i i and copepodi tes 

(Sherman and Honey 1971). 

In winter, as the larvae become 

larger (21 to 30 mm), the diet shifts 

to 1 arger fooa i terns, predominantly 

the copepods Pseudocalanus minutus and 

Oithona sp. (~herman and Honey 1971). 

-article range of sizes that 
larvae can ingest is related to the 

gape of the jaw, which increases as 

the larvae grow larger. The 

importance of a larger jaw gape, with 

an increasing size range of potential 

prey, is probably greatest during 

winter when food is scarce and small 

1 arvae of zoopl ankton are 1 ac ki ng 

(Sherman and Honey 1971). If herring 

larvae starve for 5 to 7 days, they 

essentially lose their ability to feed 

and become easy prey (Lett 1976). 

During zooplankton blooms in spring, 

herring larvae eat a wider variety of 

prey--principal ly ci rriped larvae, 

crustacean eggs, ti nti nnids, and 

copepods (Sherman and Honey 1971). 

The larval phase of Atlantic herring 

lasts 6 to 8 months (Sindermann 1979). 

Movement. Studies of 1 arval 

drift patterns in the Gulf of Maine 

indicate that larvae, soon after 

hatching , either move directly into 

estuaries and embayments adjacent to 

the spawning areas or are carried 

along the coast by east to west 

coastal currents. As an indication of 

movement by east to west currents, 

larvae hatched off eastern Maine are 

regularly collected in the Sheepscot 

Estuary of western Maine by late 

autumn (Graham and Joule 1981; Graham 

1982). The larvae make diurnal 

vertical migrations to move into the 

estuaries, using 1 andward tidal 

currents (Graham 1972). Larvae 

spawned in Georges Bank waters in 1971 

were widely dispersed but there was 

some indication of a drift toward the 

southwest. In contrast, most of those 

from spawning sites at Jeffreys Ledge, 

Cape El izabeth, and Stel lwagen Bank 

moved shoreward (Sindermann 1979). 

Juveni 1 es 

Metamorphosi s. When young 

herring are about 50 to 55 mm SL, they 

undergo metamorphosis , developing the 

morphological characteristics of 

adults; they are then identified as 

juveniles (Lough et al. 1982). 

Metamorphosis begins in April in the 

Gulf of Maine (Sindermann 1979). In 

1 aboratory studies, metamorphosis was 

completed in about 10 days at 15 OC 

(Bl axter and Hol 1 i day 1963). After 

metamorphosis, school i qg behavior 

becomes prominent and lasts throughout 

life. As young herring begin to 

school , they moved inshore in 1 arge 

numbers. 

Movement. The movements of 

juveni 1 e herring in the Cape Cod - Bay 

of Fundy area are not well known. 

Herring schools of age gr.oups I and I1 

in the Gulf of Maine are known to move 

inshore in spring and remain there 

during summer. In southwestern Maine, 

juveni les tagged during summer often 

were recovered in Massachusetts Bay 

during winter, but those tagged in 

eastern Maine in summer tended to 

remain nearby in winter (Creaser et 

al. 1984). Juveniles tagged in 

southwestern and central Maine a1 so 

were recovered farther east in the 

following summer, but most larvae from 

eastern Maine were recovered near 

where they were originally tagged. 

Fish tagged during winter in eastern 

and western Maine remained in the same 

area during the following summer 

(Creaser et a1 . 1984). 

Light response. The response of 

juvenile herring to light is strongly 

negative. The fish move toward the 

surface at sunset and remain there 

during low 1 ight intensity. 

Laboratory studies indicate that 

juveniles are most active just after 

sunset and just before sunrise 

(Stickney 1972). Commercial catches 

of juvenile herring in Maine have been 

inversely correlated with moonlight 

intensity. Monthly peaks in catches 

coincided with the dark phase of the 

moon (Anthony 1971). 

Food habits. Juveni 1 e herri ng 

feed on a variety of zooplankton. 

Copepods are the most important prey 

throughout the year (Sherman and 

Perki ns 1971). Larval decapods are 

important as food in spring and larval 

cirripeds in spring and summer. Other 

foods are larval pelecypods in summer, 
and cladocerans in summer and autumn Ann - Jeffreys Ledge area in winter 

(Sherman and Perki ns 1971). (Creaser and Libby 1982). 

Adults 

Age and maturity. In the Gulf of 

Maine and Northwest Atlantic, most 

herring of both sexes first spawn at 

age IV. Some fish become sexually 

mature at the age I11 though the 

number varies from year to year 

(Anthony and Waring 1980b). The 

percentages of Georges Bank herring 

that matured at age I11 ranged from 6 

to 62 (mean = 29) for the 1960-65 year 

classes (Boyar 1968). Length at first 

spawning averaged 26 cm TL for fish of 

age group I11 and 27.5 cm for those of 

age group IV (Boyar 1968). 

Migration. Adult herring 

sometimes travel lona distances to 

spawning areas in late summer and 

early fall, then to warmer overwintering areas after spawning. In spring 

and early summer, they undertake 

feeding migrations. Migrating herring 

schools may consist of hundreds of 

thousands of fish. Each school is 

usually composed of fish about the 

same size and age (Hi ldebrand 1963). 

They seldom migrate seaward beyond on 

depth of 50 fathoms. Herring usually 

inhabit waters closer to the surface 

than the bottom, except in midwinter 

(Hi 1 debrand 1963). 

Tagging studies have shown that 

spawners from Jeffreys Ledge may 

overwinter south of Cape Cod and 

migrate in spring and summer to 

northern or offshore areas of the Gulf 

of Maine and into the Bay of Fundy 

(New England Regional Fishery 

Management Counci 1 1978). Adul ts from 

southeast Nova Scoti a move primari ly 

northward into Chedabucto Bay in 

northeast Nova Scotia to overwinter, 

but aft.er spawning many migrate south 

and west as far as Cape Cod. Many 

adults tagged in eastern Maine in the 

summer were recovered in Massachusetts 

Bay in winter (Creaser et al. 1984). 

Fish that spawn near the New Brunswick 

coast have been reported in the Cape 

Food habits. Adult herring feed 

selectively on zooplankton, capturing 

prey by direct, predatory snapping 

action (Blaxter and Hol 1 iday 1963). 

In the North Atlantic Region, the 

principal prey is the euphausiid, 

Meganyctiphanes norvegica; of lesser 

importance are chaetognaths and the 

copepod, Cal anus f i nmarchi us (Maurer 

and ~owmanm 

Growth Characteristics 

Larval herring from coastal Maine 

grow about 2.0 mm per week from 

October to early January and from late 

February to early March; little 

growth, if any, occurs during midwinter (Townsend and Graham 1981). Other 

growth rates reported in the Gulf of 

Maine - Georges Bank area were 1.75 mm 

per week after hatching, 2.1 mm per 

week for larvae 20 days old in September and October, and (1.0 mm per week 

for fish 75 days old in winter (Lough 

et al. 1982). The average daily 

growth increment for larvae of Georges 

Bank-Nantucket Shoals was 0.2 nun 

(Lough 1976). 

Most of the growth of juvenile 

herring is in summer. In New 

Brunswick, the average growth of 

2-year-01 d herri ng duri ng May to 

September ranged from 30 mm in 1965 to 

55 mm in 1978 (Sinclair et al. 1981). 

Both sexes are about 90 to 125 mm TL 

at the end of the first year of life 

and 190 to 200 mm TL after the second 

(Bigelow and Schroeder 1953). 

Anthony (1971), who compared 

juvenile growth in different areas of 

Maine, reported that modal lengths of 

fish of the 1960 year class (age group 

11) in October in eastern, central, 

and western Maine were 142, 155, and 

175 mm, respectively. The lengths of 

the 1959 year class for the same age 

group and areas were 190, 216, and 

214 mm. In fish up to 3 year!! old, 

growth was inversely related to 
population density and was faster in 

western than in eastern Maine. 

Von Bertal anffy growth curve 

parameters, K (growth rate) and L 

(maximum length attained) , were 0.251 

and 37.4 cm in eastern Maine (Anthony 

1971). The same values for southern 

and central Maine combined were 0.267 

and 36.0 cm. A Newfoundland herring 

study provided little evidence to 

support density-dependent growth in 

the first year, but suggested 

density- i ndependent growth regulated 

by temperature (Moores and Winters 

1982). 

Anthony and Waring (1980a) 

reported that adult herring reach 

their peak weight each year in August 

or September and are lightest in 

February-March. They a1 so reported 

that herring (both sexes) from Nova 

Scotia, eastern and western Maine, and 

Jeffreys Ledge a1 1 had "oceanic" 

(offshore) growth characteristics 

common to fish of the Northeast 

Atlantic. The characteristics were 

the large maximum size, slow growth 

rate, and maximum ages of 15-18 years 

(Table 3). 

Georges Bank herring are more 

similar in growth characteristics to 

Northwest Atlantic "shelf" (inshore) 

populations, which have a smal ler 

maximum size, higher growth rate, and 

lower maximum age (Table 3; Figure 3). 

Table 3. Von Bertalanffy growth 

parameters of At1 antic herring stocks 

of Northeastern U.S. (Anthony and 

Wari ng 1980a). 

Area 

Georges Bank 34.1 0.377 13 

Jeffreys Ledge 36.7 0.311 15 

Western Maine 35.1 0.335 16 

Eastern Maine 38.0 0.277 16 

Nova Scotia 39.0 0.179 18 

Figure 3. Von Bertalanffy growth 

curve for the Georges Bank herring 

(1960-1971 year classes included). 

Fran Anthony and Waring (1980a). 

'The Von Bertalanffy growth rate (K) of 

Georges Bank herri ng i ncreased from 

0.350 (year classes 1960-63) to 0.357 

(year classes 1964-67) to 0.510 (year 

classes 1968-71). The increased 

growth rate of a1 1 year classes after 

1968 may have been related to the 

decline in abundance (Anthony and 

War i ng 1980a). 

THE FISHERY 

Hi story of the Fishery 

An extensive herring fishery 

exists between southern New England 

and Newfoundland (Figure 4). The 

At1 antic herring has supported a 

commercial fishery along inshore 

waters of New England for at least 400 

years (Anthony and Waring 1980~). The 

canning of "sardines" began in Maine 

in 1875 and, along with the sardine 

fishery of New Brunswick, continues to 

be an important economic resource. 

The coastal Maine and New Brunswick 

fisheries are primarily fixed-gear 

(stop seine and weir) fisheries that 

concentrate on juveni 1 e herri ng, 

primarily of age group I1 (Anthony and 

Waring 1980a). Currently, major 

fishing grounds in the Northwest 
Figure 4. Herring fishing areas of the Northwest Atlantic (shaded areas 

indicate major fishing grounds, though Georges Bank does not support a 

significant fishery). 

Atlantic are the Gulf of Maine, 

southwestern Nova Scotia, Chedabucto 

Bay, the Magdalen Islands area in the 

Gulf of St. Lawrence, and southwestern 

Newfound1 and. 

The juvenile catch in Maine has 

fluctuated greatly throughout the 

history of the fishery because of 

variations in recruitment, market 

demand, and vulnerability to new 

fishing gear. Yet, juveniles are 

still the dominant component of the 

Maine herring fishery (Table 4). A 

period of decline in catch began in 

the early 1950's in eastern Maine, 

historically the main source of 

juveni 1 e fish (Bigel ow and Schroeder 

1953; Sindermann 1979; Anthony and 

Waring 1980a). Declines in western 

and central Maine began in the 1960's 

and lasted through 1971. The total 

Table 4. Canmercial landings (metric 

tons) of Atlantic herring of different 

ages in Maine, 1979-82. 

Age yeara (Years) 1979 1980 1981 1982 

I 27 1,012 120 451 

I1 5,839 9,051 39,57513,311 

I11 11,485 26,538 1,627 3,410 

IV 1,708 8,606 2,586 328 

V 191 442 3,815 4,115 

V I 329 24 366 2,045 

VI I 196 101 20 194 

VIII 104 20 18 - 

>VIII 179 133 120 135 

Total 20,058 45,907 48,249 24,007 

'~ata for 1979-81 fran Chenoweth 

(1982); data for 1982 fran Fogarty 

and Clark (1983). 
In 1967, the fishery for adult 

herring in the western Gulf of Maine 

intensified, and in 1969, all fisheries combined for a harvest of 967,000 

t -- an all-time peak. Such heavy 

fishing pressure caused production to 

decline after 1973. 

catch of juveniles in 1971 was only 

5,400 t (Chenoweth 1982). The catch 

of juveniles in Maine then increased, 

peaking in 1981 at 41,200 t; however, 

the 1982 catch dropped to 17,000 t 800- 

(Fogarty and Clark 1983). 

h - 

Before 1961, only the inshore 4 

fisheries for juveniles in Maine and 

New Brunswick were important in the 

Northwest Atlantic; however, fishing 

intensity on adults by the USSR 2 

fishing fleet, primarily on Georges a 

Bank, sharply increased in 1961. 400- 

After 1961, other foreign nations also 1 

began to fish there. The Georges Bank E 

herring fishery, nonexistent in 1960, 

rose to a production of 373,600 t in 

1968 (Anthony and Waring 1980a). 200 - 

In 1964, Canadian purse seine 

vessels sharply increased their 

fishing effort in the Gulf of St. 0- 

The adult herring landings on 

Jeffreys Ledge in the western Gulf of 

Maine declined to 7,600 t in 1982 -- a 

50% drop from 1981, and a nearly 79% 

decline from the 1980 catch of 36,100 

t (Fogarty and Clark 1983). Purse 

seines and pair trawls are primarily 

used in this fishery, which is active 

during fa1 1 and winter; however, there 

is also some fishing for adults along 

the Maine coast during summer and 

early fall. Catches of adults from 

central , eastern, and western Maine 

are not as large as those from 

Jeffreys Ledge; catches in Maine in 

1978 and 1980, which were better than 

average, were 6,200 and 9,200 t, 

respectively (Chenoweth 1982). Maine 

'I 

, 

and Massachusetts contributed most of 

the New England landings in 1976-82 

(Table 1). 

Lawrence and off Newfoundland and Nova 1920 1930 1940 1950 1960 1970 

Scotia. The total catch for the Year 

Northwest Atlantic increased from Figure 5. Annual landings of Atlantic 

180,000 t in 1960 to 600,000 t in 1964 herring from the Northwest Atlantic 

(Figure 5). 1920-1975 (Anthony and Waring 1980a). 

In 1972, the International 

Commission for Northwest Atlantic 

Fisheries (ICNAF) began establishing 

total a1 lowable catches for adults. 

The catches were set more for economic 

than for conservation reasons (Anthony 

and Waring 1980a) and catches soon 

declined in all areas. The Fishery 

Conservation and Management Act of 

1976 prohibited fishing by foreign 

vessels within 200 miles of shore, an 

area which includes Georges Bank. The 

Georges Bank catch in 1977 was almost 

nil (Anthony and Waring 1980a) and has 

improved 1 i ttl e in 1978-83. 

Current management of the herring 

fishery is at the State level until 

regional management can be formulated 

and established. A management plan by 
the New England Regional Fishery 

Management Counci 1 , which establ ished 

catch quotas in Federal waters from 3 

to 200 miles offshore, was abandoned 

in 1982. 

A definition of unit stocks of 

herring in the Northwest Atlantic, 

which is essential for formulating 

management, has received much 

attention from biologists and 

managers, but has not been fully 

resol ved. Management is currently 

based on the assumption that there are 

three herring stocks or stock 

complexes from Nova Scotia to southern 

New England: Georges Bank, the Gulf 

of Maine, and southwest Nova Scotia. 

Primary criteria for separating these 

groups are differences in spawning 

population sizes, meristic counts (fin 

rays and vertebrae), and growth rates 

(Anthony 1972). Current stock assessments provide population estimates for 

both the Gulf of Maine and the combined stocks of the entire region from 

Cape Hatteras, N.C. to Nova Scotia 

(Fogarty and Clark 1983). The latter 

"pooled" assessment recognizes season 

stock intermixing throughout the Gulf 

of Maine and Nova Scotian shelf 

(Fogarty and Clark 1983). 

ECOLOGICAL ROLE 

Atlantic herring are a significant part of the diet of many predatory fishes, marine mammals, and 

piscivorous birds. They also are 

important competi tors of other planktivorous fishes, principally Atlantic 

mackerel , another commercial ly important pelagic species in the Northwest 

Atlantic. Chief predators of herring 

are cod, pol lock, haddock, si 1 ver 

hake, mackerel, dogfish and other 

sharks, finback whales, and the common 

squid (Hi 1 debrand 1963; Bigel ow and 

Schroeder 1953). School s of herring 

in the Gulf of Maine are also preyed 

upon by short-finned squid, striped 

bass, tuna, salmon, mackerel sharks 

(Bigelow and Schroeder 1953), and 

Atlantic white-sided dolphins (Katona 

et al. 1978). Herring are a common 

prey of rorqual whales (Overhol tz and 

Nicolas 1979). Sooty shearwaters feed 

on herring in eastern Canadian waters 

(Brown et a1 . 1981). 

A principal source of herring egg 

mortality is predation by ground fish, 

e. g. , cod and haddock. Hempel (1971) 

estimated that on North Sea spawning 

grounds, one haddock ate the spawn of 

one herring each day of the spawning 

season. On Georges Bank, Caddy and 

Iles (1973) observed that heavy predation on herring egg beds caused 

steep-sided craters or depressions (up 

to 0.3 m in diameter) in the egg 

1 ayer, sometimes exposing the underlying gravel. An estimated 8% of the 

spawn was removed by predators. Red 

hake were seen feeding on herring 

eggs ; and moon snai 1 s , hermit crabs, 

and starfish were found in the egg mat 

depressions (Caddy and Iles 1973). 

McKenzi e (1964) reported that haddock 

in the area off Black Point, Nova 

Scotia ate 1 arge quantities of herring 

eggs, and apparently were the main egg 

predator. On Georges Bank in fall 

1970, herring eggs made up 28% of the 

weight of haddock stomach contents 

(Langton and Bowman 1980). 

Analysis of food habits of 

Georges Bank and Gulf of Maine fishes 

in the 1960's and 19701s, in relation 

to fluctuations in herring and 

mackerel stock sizes, disclosed that 

herring accounted for 30% of the total 

weight of food eaten by Atlantic cod 

(Grosslein et al. 1980). The decline 

in herring and mackerel biomass in 

these years was accompanied by a 

decline in almost all demersal fish 

species, and crustaceans became 

slightly more important than fish in 

the cod diet (Grosslein et al. 1980). 

This study also revealed a high degree 

of diet over1 ap (71%) between herring 

and mackerel, due to the preference of 

both species for euphausiids. For 

example, euphausiids made up 51% of 

the herring diet and 52% of the 

mackerel diet on Georges Bank 

(Grosslein et a1 . 1980). 
Herring populations may be 

affected by predation of adult 

mackerel on herring larvae and by 

cannibalism of older herring on their 

own young in the Gulf of St. Lawrence 

(Lett and Kohler 1976). Apparently, 

the growth rate of yearling herring 

declined rapidly in response to an 

increase in abundance of young-ofthe-year mackerel, probably because 

both forage for zooplankton of the 

same sizes. Yearling mackerel feed on 

smaller plankton and do not compete 

heavily with herring (Lett and Kohler 

1976). 

ENVIRONMENTAL REQUIREMENTS 

Temperature 

When water temperatures are 8 to 

13 OC, the average incubation period 

of herring eggs in the Gulf of Maine 

is 10 to 15 days (Hildebrand 1963; 

Bigelow and Schroeder 1953). On 

Jeffreys Ledge, eggs hatched in 7 days 

at 13 OC and in 12 days at 8 OC (Boyar 

et al. 1973). Much of the primary 

work on temperature effects on egg and 

larval development was done with 

European herring. Meyer (1878), as 

cited by Blaxter and Holliday (1963), 

reported that egg development was 

normal at 1 to 22 OC but was fatally 

low at -0.8OC. Slightly lower 

minimum temperatures for development 

have ranged from -1.2 to 0 OC for 

herring stocks off northern Europe 

(Soleim 1942; Blaxter and Hempel 1961; 

Blaxter and Holliday 1963). The jaw 

may develop abnormally at low 

temperatures and reduce the ability of 

the larvae to capture prey (Alderdice 

and Velsen 1971). 

Two equations were developed for 

predicting hatching time (days) in 

relation to water tem erature (OC): 

D = 4 + 44.7e-O. 167f (Hela and 

Laevastu 1962); and D=-2.0 + 

165.0/T+1.34 (Bl axter 1956). 

Experiments at Grand Manan, New 

Brunswick, showed that temperatures 

above 20 OC and below 5 OC were lethal 

to eggs (Bigelow and Schroeder 1953). 

Yolk-sac stage duration varies 

with temperature. Temperatures and 

durations of the yolk-sac stage were 

2.5 days at 14.5 OC and 4.5 to 14 days 

at 8 OC (Mansueti and Hardy 1967). 

Lough et al. (1982) reported yo1 k 

absorption times of 4.5 days at 10 OC 

and 6 days at 8 OC. 

Temperature a1 so affects the 

availability of plankton for food. If 

the water temperature is too low, 

production of plankton of suitable 

size is inadequate to support an 

abundance of herring 1 arvae (Lett 

1976) ; optimal water temperatures 

induce improved growth and high 

survival of herring 1 arvae. 

Furthermore, swimming abi 1 i ty 

increases and predator avoidance and 

feeding abi 1 i ty are improved (Lett 

1976). 

Upper and 1 ower temperature 

tolerances of newly hatched herring 

larvae are 22 to 24 OC and -0.75 to 

-1.8OC for fish acclimated to 

temperatures between 7.5 and 15.5 OC 

(Blaxter and Holliday 1963). 

Barker et al. (1981) tested 

thermal to1 erances of herring under 

conditions associated with passage 

through condenser cooling systems of 

electrical generating stations. 

Larvae were exposed to rapid temperature increases from a base temperature 

of 8 OC, held for a standard period of 

time, and then rapidly returned to the 

original base temperature. The temperature changes ranged from 16 to 

25 OC at exposure times of 5, 15, 30, 

and 60 min. Larvae acclimated to 8 OC 

survived temperature changes of 17 OC 

for up to 60 min and higher temperatures (27 to 29.1 OC) for shorter (~30 

mi n) periods. 

At water temperatures of 19.5 to 

21.2 OC, mortality of juvenile herring 

(11.1 to 21.9 cm TL) was about 50% 

after 48 h. Tolerance to high 
temperatures was greater among small 

fish (Brawn 1960a). The mean freezing 

point of herring blood is -0.95 OC, 

decreasing to -1.01 OC during winter 

(Bl axter and Hol 1 i day 1963). Be1 ow 

these temperatures, herring blood 

quickly freezes unless supercool i ng or 

freezing point depression takes place 

(Blaxter and Holliday 1963). The 

freezing point is 0.75 OC for the 

fluids of ripe eggs and 0.92 OC for 

sperm and parental blood (Bl axter and 

Holliday 1963; Blaxter and Hunter 

1982). 

The migration and distribution of 

herring are linked with thermal 

oceanic fronts between colder, 1 ess 

sal i ne conti nental she1 f water and 

warmer, more sal i ne continental slope 

water (Sindermann 1979; Iles and 

Sinclair 1982). In these fronts, 

plankton and other fish food organisms 

are usually abundant. Intrusions of 

warmer slope water into spawning areas 

in the Gulf of Maine influence 

spawning success and cause annual 

variations in commercial catch 

(Si ndermann 1979). Higher water 

temperatures in September to March 

seem to favor spawning and recruitment 

success (Anthony 1972). 

Sal i ni ty 

Variations in salinity can affect 

early development, particularly of 

newly spawned eggs. Herring eggs are 

freely permeable immediately after 

spawning and, therefore, have no 

protection against osmotic imbalance 

(Holl iday 1965). Because they are 

isotonic with seawater, the eggs are 

larger than average in low salinities 

and smaller in high salinities. The 

closure of the blastopore after 

gastrulation--which occurs in 24 h at 

17 to 24 OC (Mansueti and Hardy 

1967)--causes the egg to become more 

tolerant to changes in salinity. In 

general , both extremes of sal i ni ty are 

damaging (Bl axter 1965). 

Laboratory studies indicate that 

ferti 1 ization, egg development, and 

hatching can succeed in salinities of 

5.9 to 52.5 ppt (Holliday and Blaxter 

1960). Maximum fertilization is at 25 

ppt or more, and hatching success is 

greatest at 20 to 35 ppt. Egg 

ferti 1 izations reported by Holl iday 

and Blaxter (1961) were 70% at 5 to 12 

ppt, and 100% at 25 to 55 ppt. 

Atlantic herring larvae tolerate 

a wide range of salinities under 

experimental conditions. Bl axter and 

Hunter (1982) reported a tolerance to 

salinities of 1.4 to 60 ppt for 24 h 

and 2.5 to 52.5 ppt for 7 days. The 

salinity isosmotic with body fluids 

was 12 ppt. Yolk-sac larvae, however, 

survive longer in salinities between 

10 and 20 ppt (Holliday 1965). The 

plasticity that is apparent in 

salinity tolerance of Atlantic herring 

eggs and larvae may be indicative of 

divergent physiological races 

(Alderdice et al. 1979). 

Atlantic herring show an apparent 

preference for higher salinities as 

they become older. The lower level of 

tolerance for juveniles is about 5 ppt 

(Brawn 1960b). A1 though herring enter 

Northwest Atlantic bays and estuaries 

freely, they are rarely observed in 

salinities less than about 3 ppt 

(Hi 1 debrand 1963). 

A1 though Atlantic herring in 

European waters sometimes spawn in 

shallow inshore waters with salinities 

of 5 to 35 ppt (Alderdice et al. 

1979), herring from the Northwest 

Atlantic are known to spawn only in 

water of 32 to 33 ppt (Hildebrand 

1963). 

Oxygen 

Probably because oxygen is rarely 

a limiting factor in the marine 

environment, studies on oxygen-related 

effects on herring are scarce. 

DeSilva and Tytler (1973, cited in 

Blaxter and Hunter 1982) reported that 

larval 96 h LDS0 at 10 OC ranged from 

about 1.9 to 3.6 mg/l. Braum (1973) 

reported low hatching success for 
herring eggs held in water with 

dissolved oxygen be1 ow 20% saturation. 

Larvae 3 to 4 days old died in 6 h at 

11.6% oxygen saturation (Bishai 1960). 

Sediments and Turbidity 

A proposal to dredge a channel 35 

nautical miles long in Miramichi Bay, 

New Brunswick, an important herring 

fishing ground, prompted Messieh et 

al. (1981) to investigate the effects 

of suspended silt and clay sediments 

on early life stages of Atlantic 

herring. Normal suspended sediment 

concentrations of less than 20 mg/l 

were expected to be replaced by 

concentrations of up to 2,000 mg/l 

duri ng dredging and dumping. 

Experiments conducted by Messieh et 

al. found 100% mortality in eggs 

covered with 1 cm of sediment and 85% 

mortality in eyed eggs covered with 

only a thin film of sediment. These 

tests indicate that at least a portion 

of the egg must be exposed free of 

sediment if the egg is to survive. 

Though hatching success apparently was not affected by suspended sediment concentration or egg density, 

larvae hatching at 1 ow sediment concentrations (0-540 mg/l) tended to be 

larger. High egg density tends to 

cause premature hatchi ng and 1 arvae at 

hatching were smallest when egg 

densities were high, regardless of 

sediment concentration. Messieh et 

al. (1981) noted no deleterious 

effects of suspended sediments on 

hatching success at any sediment 

concentrations up to 7,000 mg/l. 

Hatched 1 arvae suffered 100% mortal i ty 

after 48 h at 19,000 mg/l. 

Tests indicated that juveni 1 e 

At1 antic herring avoided suspended 

sediment concentrations between 9.5 

and 12 mg/l; some juveniles avoided 

concentrations as low as 2.5 mg/l. In 

feeding experiments, the number of 

larval herring feeding at any one time 

was significantly reduced at suspended 

sediment concentrations greater than 

3.0 mg/l (Messieh et al. 1981). 

Water Movement 

Water currents are important to 

Atlantic herring in the Gulf of Maine 

and on Georges Bank because they 

transport and entrain 1 arvae and 

plankton into estuaries and coastal 

areas during autumn and winter. The 

net a1 ongshore drift carrying 1 arvae 

in the Gulf of Maine is principally 

shoreward from east to west (Graham 

1970). Larvae mi grate vertical ly in 

the water column to take advantage of 

landward tidal currents (Graham 1972). 

Adequate water exchange is an 

important environmental requirement 

for herring spawning grounds. 

Sindermann (1979) reported that a 

current velocity of at least 0.27 to 

0.52 m/sec must be present; Caddy and 

Iles (1973) observed bottom currents 

of 0.5 to 1.0 m/sec on a Georges Bank 

spawni ng area. 

Environmental Contaminants 

The effects of copper on eggs and 

1 arvae of At1 antic herring were 

reported by Bl axter (1977). Mortal i ty 

of newly hatched larvae was high at 

copper concentrations of 1,000 

micrograms per 1 i ter. Eggs incubated 

in 30 1 during incubation had 

relatively high mortality and 

premature hatching; 70% of the larvae 

hatched were deformed. Larvae were 

more resistant to copper than eggs; 

survival of larvae was impaired only 

at concentrations of > 1,000 pg/l. 

The vertical migration -of larvae was 

impaired at copper concentrations of - > 

300 pg/l. 

Tests on the effects of sulfuric 

pol 1 utants (FeSO, and H2S04) showed 

that a dilution of 1:8,000 significantly reduced egg ferti 1 ization and 

hatching success, decreased egg di ameter, retarded embryonic growth, 

shortened the incubation period, and 

increased the rate of structural 

abnormal i ties in newly hatched 1 arvae 

(Ki nne and Rosenthal 1967). Larval 

prey-catchi ng abi 1 i ty was impai red in 
1:32,000 and 1:24,000 dilutions; 

1 ocomotory performance was seriously 

affected at a 1:16,000 dilution; and 

paralysis, shrinkage, permanent 

deformities, and death occurred within 

a few days at 1:8,000 dilution. 

Studies of di ni trophenol effects 

on herring embryonic development 

indicated that low concentrations 

(0.01 to 0.05 pmol/l) increased embryo 

activity and altered heart rates 

significantly (Rosenthal and Stelzer 

1970). Various embryonic 

ma1 formations were a1 so observed. A 

dinitrophenol concentration of 0.1 

pmol/l caused an up to 400% increase 

in the normal embryonic respiration 

rate (Stelzer et a1 . 1971). 

Bl axter and Hunter (1982) 

reported that eggs and larvae held 

under films of crude oil in 

concentrations of 1 to 20 ml/l , or in 

emulsions, experienced toxicities that 

varied with the origin of the oil. 

For oil from a particular source, the 

fractions with the lower boiling 

points seemed more harmful (Kuhnhold 

1969). In tests on oil dispersants,, 

larvae did not avoid horizontal 

gradients, but swam into surface 

dispersant 1 ayers and were narcotized 

(Wilson 1974). The survival of! 

herring eggs and larvae was highest ip 

water with low biological oxygej 

demand and low nitrate levels (Baxte 

and Steel e 1973). 
LITERATURE CITED 

A1 derdice, D. F. , and F. P. J. Vel sen. 

1971. Some effects of salinity and 

temperature on early development of 

~aci fc herring (~l upea dal 1 asi ). 

J. Fish. Res. Board Can. 

Alderdice, D.F., T.R. Rao, and H. 

Rosenthal. 1979. Osmotic responses 

of eggs and larvae of the Pacific 

herring to salinity and cadmium. 

Helgol . Wiss. Meeresunters. 32: 508- 

538. 

American Fisheries Society , Committee 

on Names of Fishes. 1980. A list 

of common and scientific names of 

fishes from the United States and 

Canada, 4th ed. Am. Fish. Soc. Spec. 

Publ. No. 12. 174pp. 

Anthony, V.C. 1971. The density 

dependence of growth of the Atlantic 

herring in Maine. Rapp. P.-V. Reun. 

Cons. Int. Explor. Mer 160:197-205. 

Anthony, V. C. 1972. Population 

dynamics of the Atlantic herring in 

the Gulf of Maine. Ph.D. Thesis. 

University of Washington, Seattle. 

266 pp. 

Anthony, V.C. , and G.T. Waring. 

1980a. The assessment and 

management of the Georges Bank 

herring fishery. Rapp. P. -V. Reun. 

Cons. Int. Expl or. Mer 177: 72-111. 

Anthony, V.C., and G.T. Waring. 

1980b. Estimates of herring 

spawning stock biomass and egg 

production for the Georges Bank - 

Gulf of Maine region. Northwest 

Atl. Fish. Organ. SCR Doc. 

80/1X/135. 

Anthony, V.C. , and G.T. Waring. 

1980c. A review of the herring 

fisheries, their assessment, and 

management in the Georges Bank - 

Gulf of Maine area. Pages 115-178 

- in Proceedings of the Alaska herring 

sympos i um, February 19-21, 1980. 

Anchorage, A1 as ka. 

Barker, S. L. , D. W. Townsend, and J. S. 

Hacunda. 1981. Mortal i ties of 

Atlantic herring, Clupea h. 

harengus , smooth f 1 ounder, ~iopsetTa 

putnami , and rainbow smelt , Osmerus 

mordax larvae exposed to acute -9 

thermal shock. U.S. Natl. Mar. 

Fish. Serv. Fish. Bull. 

79(1): 198-200. 

Baxter, I. G. 1971. Development rates 

and mortalities in Clyde herring. 

Rapp. P.-V. Reun. Cons. Int. Explor. 

Mer 160: 27-29. 

Baxter, I.G. , and J.H. Steele. 1973. 

Mortality of herring larvae in the 

Clyde Sea area. Int. Counc. Explor. 

Sea, Fish. Improv. Comm. Pap. E29. 

7 pp. (mimeo.) 

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

1953. Fishes of the Gulf of Maine. 

U.S. Fish Wildl. Serv. Fish Bull. 

53. 577 pp. 

Bishai, H.M. 1960. The effect of gas 

content of water on larvae and young 

fish. 2. Wiss. 2001. 163:37-64. 

Blaxter, J.H. S. 1956. Herri ng 

rearing 11. The effect of 

temperature and other factors on 

development. Department of 

Agriculture and Fisheries for 

Scotland, Mar. Res. No. 5. 19 pp. 
Blaxter, J.H.S. 1965. The feeding of 

herring larvae and their ecology in 

relation to feeding. Calif. Coop. 

Oceanic Fish. Invest. Rep. 10: 79-88. 

Blaxter, J.H.S. 1977. The effect of 

copper on the eggs and larvae of 

plaice and herring. J. Mar. Biol. 

Assoc. U. K. 57:849-858. 

Bl axter, J. H. S. , and G. Hempel. 1961. 

Biologische Beobachtungen bei der 

Aufzucht von Heringsbrut. Helgol. 

Wiss. Meeresunters. 7: 260-283. 

Blaxter, J.H.S., and F.G.T. Holliday. 

1963. The behavior and physiology 

of herring and other clupeids. Adv. 

Mar. Biol . 1:261-393. 

Bl axter, J. H. S. , and D. E. Hoss. 1979. 

The effect of rapid changes of 

hydrostatic pressure on the Atlantic 

herring Clupea harengus L. 11. The 

response of the auditory bull a 

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1960. Histological changes in the 

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Graham, J.J. 1972. Retention of 

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160: 8-11. 

Hempel, G., and K. Schubert. 1969. 

Sterbl inchkeitsbestimmug an einem 

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U. K. 39: 591-603. 

Holliday, F.G.T., and J.H.S. Blaxter. 

1961. The effects of salinity on 

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Mar. Biol. Assoc. U.K. 41:37-48. 

Hoss, D. E. , and J. H. S. Bl axter. 1979. 

The effect of rapid changes of 

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Iles, T.D., and M. Sinclair. 1982. 

At1 antic herring: stock di screteness and abundance. Science 215: 

627-633. 

Katona, S.K., S.A. Testaverde, and B. 

Barr. 1978. Observations on whitesided dolphin, La enorh nchus 

acutus, probably killed + in gi 1 nets 

in the Gulf of Maine. U.S. Natl. 
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475-476. 

Kelly, K.H. 1983. Comparison of 

reproductive characteristics and age 

composition of spawning groups of 

Atlantic herring in the Gulf of 

Maine. M.S. Thesis. University of 

Maine, Orono. 99 pp. 

Kinne, 0. , and H. Rosenthal . 1967. 

Effects of sulfuric water pollutants 

on fertilization, embryonic 

develo~ment and larvae of the 

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(Berl . ) 1: 65-83. 

Kuhnhold, W.W. 1969. Der Einfluss 

wasserloslicher Bestandteile von 

Roholen und Rohol fraktionen auf die 

Entiwickelung von Heringsbrut. Ber. 

Dtsch. Wiss. Komm. Meeresforsch. 

20: 165-171. 

Langton, R. W. , and R. E. Bowman. 1980. 

Food of fifteen northwest At1 antic 

gadiform fishes. NOAA (Natl . Ocean. 

Atmos. Adm. ) Tech. Rep. NMFS (Natl. 

Mar. Fish. Serv.) SSRF (Spec. Sci. 

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Lett, P.F. 1976. A review of 

density-dependent and independent 

processes which may affect 

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Intl. Comm. Northwest Atl. Fish. 

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Lett, P. F. , and A.C. Kohler. 1976. 

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Intl. Comm. Northwest Atl. Fish. 

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PP. 

Lough, R.G. 1976. Analysis of various 

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Abundance of sea herring (Clupea 

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humpback whale saptera 

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development of herring Cl u ea 

d haren us. Mar. Biol. (Berl.)* 

Sherman, K., and K.A. Honey. 1971. 

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juveni 1e-herring fishery. Proc. 

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changes --%- in i umination. Ecoloclv 

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Pub1 ished for National Science 

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herring (Clupea harengus L. ) in 

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McMul lon. 1963. An investigation 

of herring (Cl upea harengus L. ) 

spawni na usi na free-divi na 

techniqies. 3. Fish. Res. Board 

Can. 20: 1067-1079. 

Townsend, D.W., and 3.3. Graham. 

1981. Growth and age structure of 

larval Atlantic herring, Cl upea 

haren us harengus, in the Sheepscot 

dstuary , Maine, as determined 

by daily growth increments in 

otoliths. U.S. Natl. Mar. Fish. 

Serv. Fish. Bull. 79(1):123-130. 

Wilson, K.W. 1974. The ability of 

herring and plaice larvae to avoid 

concentrations of oi 1 dispersants. 

Pages 589-602 in J.H.S. Blaxter, ed. 

The early lit history of fish. 

Spri nger-Verl ag, Berl in. 

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efficiency of spawning of Atlantic 

herrings in the Gulf of Maine. 

Polar Scientific Research Institute 

of Marine Fisheries and Oceanography 

17: 249-262. 
Estuaries Life- cycles Fisheries 

Fishes Temperature Sediment 

Growth Salinity Contaminants 

Feeding habits 

#.ntM.mlOpMtnd.d 7- 

At1 antic herring 

Cl upea harengus Spawning 

Habitat requirements 

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Unl imi ted ~nclassiGd 2 2 

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-kc. 

Aprll 1986 

a 

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

.I& Ak(- furmt: )QO rab) 

Species profiles are 1 i terature summaries ?f the morphology, range, 1 ife history, 

and environmental requirements of coastal aquatic species of animals. They are 

designed to assist in environmental impact assessment. Atlantic herring 

Clu ea haren us) spawn offshore, depositing sheets of eggs on the bottom. They 

usua 9r-f- y spawn n late summer or in fall. Fecundity of Atlantic herring in the 

Gulf of Maine ranged from 33,000 to 185,000 eggsifemale. After spawning, adults 

usually migrate to warmer areas to overwinter. All life stages feed on 

zooplankton. Adult herring reach their peak weight in August or September. 'The 

Atlantic herring supports a major fishery in Massachusetts and Maine; the 

average annual value of Maine landings was $4.3 million for 1976-82. Atlantic 

herring are significant in the diet of many predators. For juvenile herring, the 

lower level of salinity tolerance is about 5 ppt; older herring probably prefer 

higher salinities, and spawning occurs only at 32-33 ppt. Sediment deposition 

may cause egg mortality. Copper concentrations of ,1,000 mgll and sulfuric acid 

and ferrous sulfate at dilutions of L1:8,000 cause heavy larval mortality. 

REPORT WUY ENTATION ! @cmrr *- 

Pffit 1 Bioloqical Re~ort 82(11.38)* 

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

a 

*namd&u*c. 

Species Profiles: Life Histories and Environmental 

Requirements of Coastal Fishes and Invertebrates (North 

Atlantic)--Atlantic Herrinq 

7. &~tbrf.) 

Kevin H. Kelly and John R. Moring 

h -1- ~lynlr- -ma a~ wMaine Cooperative Fish and Wild1 ife Research Unit 

313 Murray Hall 

University of Maine 

Orono, ME 04469 

U~~Olynimlm)(.nwdMdnn (Q, 

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

Division of Biological Services Waterways Experiment Station 

Fish and Wildlife Service P.O. Box 631 

U.S. Department of the Interior Vicksburg, MS 39180 

Washington, DC 20240 
REGION 1 

Regional Director 

U.S. Fish and Wildlife Service 

Lloyd Five Hundred Building, Suite 1692 

500 N.E. Multnornah 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 87 103 Twin Cities, Minnesota 55 1 1 1 

REGION 5 REGION 6 

Regional Director Regional Director 

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

One Gateway Center P.O. Box 25486 

Newton Corner, Massachusetts 02158 Denver Federal Center 

Denver, Colorado 80225 

REGION 7 

Regional Director 

U.S. Fish and Wildlife Service 

101 1 E. Tudor Road 

Anchorage, Alaska 99503 
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