NOAA Technical Memorandum NMFS-NE-126

Essential Fish Habitat Source Document:

Atlantic Herring, Clupea harengus,

Life History and Habitat Characteristics

U. S. DEPARTMENT OF COMMERCE

National Oceanic and Atmospheric Administration

National Marine Fisheries Service

Northeast Region

Northeast Fisheries Science Center

Woods Hole, Massachusetts

September 1999
Recent Issues

105. Review of American Lobster (Homarus americanus) Habitat Requirements and Responses to Contaminant Exposures.

By Renee Mercaldo-Allen and Catherine A. Kuropat. July 1994. v + 52 p., 29 tables. NTIS Access. No. PB96-115555.

106. Selected Living Resources, Habitat Conditions, and Human Perturbations of the Gulf of Maine: Environmental and

Ecological Considerations for Fishery Management. By Richard W. Langton, John B. Pearce, and Jon A. Gibson, eds.

August 1994. iv + 70 p., 2 figs., 6 tables. NTIS Access. No. PB95-270906.

107. Invertebrate Neoplasia: Initiation and Promotion Mechanisms -- Proceedings of an International Workshop, 23 June

1992, Washington, D.C. By A. Rosenfield, F.G. Kern, and B.J. Keller, comps. & eds. September 1994. v + 31 p., 8 figs.,

3 tables. NTIS Access. No. PB96-164801.

108. Status of Fishery Resources off the Northeastern United States for 1994. By Conservation and Utilization Division,

Northeast Fisheries Science Center. January 1995. iv + 140 p., 71 figs., 75 tables. NTIS Access. No. PB95-263414.

109. Proceedings of the Symposium on the Potential for Development of Aquaculture in Massachusetts: 15-17 February 1995,

Chatham/Edgartown/Dartmouth, Massachusetts. By Carlos A. Castro and Scott J. Soares, comps. & eds. January 1996.

v + 26 p., 1 fig., 2 tables. NTIS Access. No. PB97-103782.

110. Length-Length and Length-Weight Relationships for 13 Shark Species from the Western North Atlantic. By Nancy E.

Kohler, John G. Casey, Patricia A. Turner. May 1996. iv + 22 p., 4 figs., 15 tables. NTIS Access. No. PB97-135032.

111. Review and Evaluation of the 1994 Experimental Fishery in Closed Area II on Georges Bank. By Patricia A. Gerrior,

Fredric M. Serchuk, Kathleen C. Mays, John F. Kenney, and Peter D. Colosi. October 1996. v + 52 p., 24 figs., 20 tables. NTIS

Access. No. PB98-119159.

112. Data Description and Statistical Summary of the 1983-92 Cost-Earnings Data Base for Northeast U.S. Commercial

Fishing Vessels: A Guide to Understanding and Use of the Data Base. By Amy B. Gautam and Andrew W. Kitts. December

1996. v + 21 p., 11 figs., 14 tables. NTIS Access. No. PB97-169320.

113. Individual Vessel Behavior in the Northeast Otter Trawl Fleet during 1982-92. By Barbara Pollard Rountree. August 1997.

v + 50 p., 1 fig., 40 tables. NTIS Access. No. PB99-169997.

114. U.S. Atlantic and Gulf of Mexico Marine Mammal Stock Assessments -- 1996. By Gordon T. Waring, Debra L. Palka, Keith

D. Mullin, James H.W. Hain, Larry J. Hansen, and Kathryn D. Bisack. October 1997. viii + 250 p., 42 figs., 47 tables. NTIS

Access. No. PB98-112345.

115. Status of Fishery Resources off the Northeastern United States for 1998. By Stephen H. Clark, ed. September 1998. vi

+ 149 p., 70 figs., 80 tables. NTIS Access. No. PB99-129694.

116. U.S. Atlantic Marine Mammal Stock Assessments -- 1998. By Gordon T. Waring, Debra L. Palka, Phillip J. Clapham, Steven

Swartz, Marjorie C. Rossman, Timothy V.N. Cole, Kathryn D. Bisack, and Larry J. Hansen. February 1999. vii + 182 p., 16

figs., 56 tables. NTIS Access. No. PB99-134140.

117. Review of Distribution of the Long-finned Pilot Whale (Globicephala melas) in the North Atlantic and Mediterranean.

By Alan A. Abend and Tim D. Smith. April 1999. vi + 22 p., 14 figs., 3 tables. NTIS Access. No. PB99-165029.

118. Tautog (Tautoga onitis) Life History and Habitat Requirements. By Frank W. Steimle and Patricia A. Shaheen. May 1999.

vi + 23 p., 1 fig., 1 table. NTIS Access. No. PB99-165011.

119. Data Needs for Economic Analysis of Fishery Management Regulations. By Andrew W. Kitts and Scott R. Steinback.

August 1999. iv + 48 p., 10 figs., 22 tables. NTIS Access. No. PB99-171456.

120. Marine Mammal Research Program of the Northeast Fisheries Science Center during 1990-95. By Janeen M. Quintal and

Tim D. Smith. September 1999. v + 28 p., 4 tables, 4 app. NTIS Access. No. PB2000-100809.
U. S. DEPARTMENT OF COMMERCE

William Daley, Secretary

National Oceanic and Atmospheric Administration

D. James Baker, Administrator

National Marine Fisheries Service

Penelope D. Dalton, Assistant Administrator for Fisheries

Northeast Region

Northeast Fisheries Science Center

Woods Hole, Massachusetts

September 1999

Essential Fish Habitat Source Document:

Atlantic Herring, Clupea harengus,

Life History and Habitat Characteristics

Robert N. Reid, Luca M. Cargnelli, Sara J. Griesbach, David B. Packer,

Donna L. Johnson, Christine A. Zetlin, Wallace W. Morse,

and Peter L. Berrien

National Marine Fisheries Serv., James J. Howard Marine Sciences Lab., 74 Magruder Rd., Highlands, NJ 07732

This series represents a secondary level of scientifiic publishing. All issues employ

thorough internal scientific review; some issues employ external scientific review.

Reviews are -- by design -- transparent collegial reviews, not anonymous peer reviews.

All issues may be cited in formal scientific communications.

NOAA Technical Memorandum NMFS-NE-126
Editorial Notes on Issues 122-152

in the

NOAA Technical Memorandum NMFS-NE Series

Editorial Production

For Issues 122-152, staff of the Northeast Fisheries Science Center's (NEFSC's) Ecosystems Processes Division have

largely assumed the role of staff of the NEFSC's Editorial Office for technical and copy editing, type composition, and

page layout. Other than the four covers (inside and outside, front and back) and first two preliminary pages, all preprinting

editorial production has been performed by, and all credit for such production rightfully belongs to, the authors and

acknowledgees of each issue, as well as those noted below in "Special Acknowledgments."

Special Acknowledgments

David B. Packer, Sara J. Griesbach, and Luca M. Cargnelli coordinated virtually all aspects of the preprinting editorial

production, as well as performed virtually all technical and copy editing, type composition, and page layout, of Issues

122-152. Rande R. Cross, Claire L. Steimle, and Judy D. Berrien conducted the literature searching, citation checking,

and bibliographic styling for Issues 122-152. Joseph J. Vitaliano produced all of the food habits figures in Issues 122-

152.

Internet Availability

Issues 122-152 are being copublished, i.e., both as paper copies and as web postings. All web postings are, or will soon

be, available at: www.nefsc.nmfs.gov/nefsc/habitat/efh. Also, all web postings will be in "PDF" format.

Information Updating

By federal regulation, all information specific to Issues 122-152 must be updated at least every five years. All official

updates will appear in the web postings. Paper copies will be reissued only when and if new information associated with

Issues 122-152 is significant enough to warrant a reprinting of a given issue. All updated and/or reprinted issues will retain

the original issue number, but bear a "Revised (Month Year)" label.

Species Names

The NMFS Northeast Regions policy on the use of species names in all technical communications is generally to follow

the American Fisheries Societys lists of scientific and common names for fishes (i.e., Robins et al. 1991a), mollusks (i.e.,

Turgeon et al. 1998b

), and decapod crustaceans (i.e., Williams et al. 1989c), and to follow the Society for Marine

Mammalogy's guidance on scientific and common names for marine mammals (i.e., Rice 1998d). Exceptions to this policy

occur when there are subsequent compelling revisions in the classifications of species, resulting in changes in the names

of species (e.g., Cooper and Chapleau 1998e

).

a

Robins, C.R. (chair); Bailey, R.M.; Bond, C.E.; Brooker, J.R.; Lachner, E.A.; Lea, R.N.; Scott, W.B. 1991. Common and scientific names of fishes

from the United States and Canada. 5th ed. Amer. Fish. Soc. Spec. Publ. 20; 183 p.

b

Turgeon, D.D. (chair); Quinn, J.F., Jr.; Bogan, A.E.; Coan, E.V.; Hochberg, F.G.; Lyons, W.G.; Mikkelsen, P.M.; Neves, R.J.; Roper, C.F.E.;

Rosenberg, G.; Roth, B.; Scheltema, A.; Thompson, F.G.; Vecchione, M.; Williams, J.D. 1998. Common and scientific names of aquatic

invertebrates from the United States and Canada: mollusks. 2nd ed. Amer. Fish. Soc. Spec. Publ. 26; 526 p.

c

Williams, A.B. (chair); Abele, L.G.; Felder, D.L.; Hobbs, H.H., Jr.; Manning, R.B.; McLaughlin, P.A.; Pérez Farfante, I. 1989. Common and

scientific names of aquatic invertebrates from the United States and Canada: decapod crustaceans. Amer. Fish. Soc. Spec. Publ. 17; 77 p.

d

Rice, D.W. 1998. Marine mammals of the world: systematics and distribution. Soc. Mar. Mammal. Spec. Publ. 4; 231 p.

e

Cooper, J.A.; Chapleau, F. 1998. Monophyly and interrelationships of the family Pleuronectidae (Pleuronectiformes), with a revised classification.

Fish. Bull. (U.S.) 96:686-726.
Page iii

FOREWORD

One of the greatest long-term threats to the viability of

commercial and recreational fisheries is the continuing

loss of marine, estuarine, and other aquatic habitats.

Magnuson-Stevens Fishery Conservation and

Management Act (October 11, 1996)

The long-term viability of living marine resources

depends on protection of their habitat.

NMFS Strategic Plan for Fisheries

Research (February 1998)

The Magnuson-Stevens Fishery Conservation and

Management Act (MSFCMA), which was reauthorized

and amended by the Sustainable Fisheries Act (1996),

requires the eight regional fishery management councils to

describe and identify essential fish habitat (EFH) in their

respective regions, to specify actions to conserve and

enhance that EFH, and to minimize the adverse effects of

fishing on EFH. Congress defined EFH as “those waters

and substrate necessary to fish for spawning, breeding,

feeding or growth to maturity.” The MSFCMA requires

NMFS to assist the regional fishery management councils

in the implementation of EFH in their respective fishery

management plans.

NMFS has taken a broad view of habitat as the area

used by fish throughout their life cycle. Fish use habitat

for spawning, feeding, nursery, migration, and shelter, but

most habitats provide only a subset of these functions.

Fish may change habitats with changes in life history

stage, seasonal and geographic distributions, abundance,

and interactions with other species. The type of habitat,

as well as its attributes and functions, are important for

sustaining the production of managed species.

The Northeast Fisheries Science Center compiled the

available information on the distribution, abundance, and

habitat requirements for each of the species managed by

the New England and Mid-Atlantic Fishery Management

Councils. That information is presented in this series of

30 EFH species reports (plus one consolidated methods

report). The EFH species reports comprise a survey of the

important literature as well as original analyses of fisheryJAMES J. HOWARD MARINE SCIENCES LABORATORY

HIGHLANDS, NEW JERSEY

SEPTEMBER 1999

independent data sets from NMFS and several coastal

states. The species reports are also the source for the

current EFH designations by the New England and MidAtlantic Fishery Management Councils, and have

understandably begun to be referred to as the “EFH source

documents.”

NMFS provided guidance to the regional fishery

management councils for identifying and describing EFH

of their managed species. Consistent with this guidance,

the species reports present information on current and

historic stock sizes, geographic range, and the period and

location of major life history stages. The habitats of

managed species are described by the physical, chemical,

and biological components of the ecosystem where the

species occur. Information on the habitat requirements is

provided for each life history stage, and it includes, where

available, habitat and environmental variables that control

or limit distribution, abundance, growth, reproduction,

mortality, and productivity.

Identifying and describing EFH are the first steps in

the process of protecting, conserving, and enhancing

essential habitats of the managed species. Ultimately,

NMFS, the regional fishery management councils, fishing

participants, Federal and state agencies, and other

organizations will have to cooperate to achieve the habitat

goals established by the MSFCMA.

A historical note: the EFH species reports effectively

recommence a series of reports published by the NMFS

Sandy Hook (New Jersey) Laboratory (now formally

known as the James J. Howard Marine Sciences

Laboratory) from 1977 to 1982. These reports, which

were formally labeled as Sandy Hook Laboratory

Technical Series Reports, but informally known as “Sandy

Hook Bluebooks,” summarized biological and fisheries

data for 18 economically important species. The fact that

the bluebooks continue to be used two decades after their

publication persuaded us to make their successors – the 30

EFH source documents – available to the public through

publication in the NOAA Technical Memorandum NMFSNE series.

JEFFREY N. CROSS, CHIEF

ECOSYSTEMS PROCESSES DIVISION

NORTHEAST FISHERIES SCIENCE CENTER
Page v

Contents

Introduction...............................................................................................................................................................................................1

Life History...............................................................................................................................................................................................1

Habitat Characteristics ..............................................................................................................................................................................4

Geographical Distribution .........................................................................................................................................................................5

Status of the Stocks ...................................................................................................................................................................................7

Research Needs .........................................................................................................................................................................................7

Acknowledgments.....................................................................................................................................................................................7

References Cited .......................................................................................................................................................................................7

Tables

Table 1. Size and age at maturity of Atlantic herring, Clupea harengus................................................................................................12

Table 2. Summary of life history and habitat parameters for Atlantic herring, Clupea harengus - Georges Bank ................................13

Table 3. Summary of life history and habitat parameters for Atlantic herring, Clupea harengus - Coastal Gulf of Maine ...................15

Table 4. Summary of life history and habitat parameters for Atlantic herring, Clupea harengus - Nova Scotia ...................................17

Table 5. Summary of life history and habitat parameters for Atlantic herring, Clupea harengus - No specific location given .............18

Table 6. Relative abundance of Atlantic herring (Clupea harengus) in New England and Mid-Atlantic estuaries by salinity zone......20

Figures

Figure 1. The Atlantic herring, Clupea harengus L. (from Goode 1884). ..........................................................................................22

Figure 2. Location of Atlantic herring spawning populations within the Gulf of Maine area.............................................................23

Figure 3. Principal spawning grounds on Georges Bank, 1964-1971, with a comparison of egg patch sizes among years................24

Figure 4. Abundance of the major prey items in the diet of Atlantic herring based on NEFSC trawl survey data on food habits......25

Figure 5. Hypothesized seasonal movements of three Atlantic herring spawning stocks inhabiting U.S. waters...............................26

Figure 6. Water column temperature and depth at stations where Atlantic herring larvae were collected during NEFSC surveys ....27

Figure 7a. Bottom water temperature and depth at stations where juveniles were collected during NEFSC bottom trawl surveys .....28

Figure 7b. Bottom water temperature and depth at stations where adults were collected during NEFSC bottom trawl surveys ..........29

Figure 8. Water temperature and depth at stations where juveniles and adults were collected during Massachusetts trawl surveys..30

Figure 9a. Water temperature and depth at stations where juveniles were collected during Rhode Island Narragansett Bay surveys .31

Figure 9b. Water temperature and depth at stations where adults were collected during Rhode Island Narragansett Bay surveys ......32

Figure 10a. Water temperature, depth, DO, and salinity at stations where juveniles were collected during Hudson-Raritan surveys ...33

Figure 10b. Water temperature, depth, DO, and salinity at stations where adults were collected during Hudson-Raritan surveys ........34

Figure 11. Distribution of Atlantic herring larvae collected during NEFSC MARMAP ichthyoplankton surveys...............................35

Figure 12. Changes in abundance of Atlantic herring larvae from 1971-1990 .....................................................................................39

Figure 13. Distribution and abundance of juvenile and adult Atlantic herring collected during NEFSC bottom trawl surveys ...........40

Figure 14. Distribution and abundance of juvenile and adult Atlantic herring collected during Massachusetts trawl surveys.............42

Figure 15. Distribution and abundance of juvenile and adult Atlantic herring collected during Rhode Island bottom trawl surveys...43

Figure 16. Distribution and abundance of Atlantic herring collected in Long Island Sound during Connecticut trawl surveys...........45

Figure 17. Distribution and abundance of juvenile and adult Atlantic herring collected during Hudson-Raritan trawl surveys...........46

Figure 18. Commercial landings and estimated stock biomass index (ages 2+) of Atlantic herring, 1967-1992..................................48
Page 1

INTRODUCTION

The Atlantic herring, Clupea harengus (Figure 1), is

a schooling, coastal pelagic species that inhabits both

sides of the North Atlantic Ocean. In the western North

Atlantic they range from Labrador to Cape Hatteras where

spring and autumn spawning populations support major

commercial fisheries (Messieh 1988). Juveniles and

adults undergo complex north-south and inshore-offshore

migrations for feeding, spawning, and overwintering. In

U.S. waters, herring from the Gulf of Maine and Georges

Bank are assessed and managed as a single stock complex

with two major spawning components (Atlantic States

Marine Fisheries Commission 1995; Northeast Fisheries

Science Center 1996).

This report provides information on the life history

and habitat characteristics of Atlantic herring stocks that

inhabit U.S. waters. This includes spawning populations

of the Gulf of Maine and Georges Bank/Nantucket

Shoals, as well as the southwestern Nova Scotia

population that is believed to mix with the coastal Maine

herring population (Stobo 1983).

LIFE HISTORY

This section provides a brief review of the biology of

Gulf of Maine area Atlantic herring. More detailed

reviews are provided by Bigelow and Schroeder (1953),

Sindermann (1979), Kelly and Moring 1986, the Atlantic

States Marine Fisheries Commission (1998), Munroe (in

prep.), and Tupper et al. (in prep.).

EGGS

Herring deposit demersal eggs on a variety of

substrates ranging from boulders, rocks, and gravel to

sand, shell fragments, and macrophytes in 20 to 80 m of

water in areas with strong tidal currents. The eggs are

1.0-1.4 mm in diameter (Bigelow and Schroeder 1953;

Fahay 1983) and adhere to the bottom, forming extensive

egg beds that are often many layers deep (Stevenson and

Knowles 1988). Gravel is the preferred spawning

substrate (Drapeau 1973), but eggs have been reported on

aquatic macrophytes on Jeffreys Ledge (Cooper et al.

1975) and in the Gulf of St. Lawrence, (Messieh et al.

1985). The eggs hatch in 10-15 days (Bigelow and

Schroeder 1953).

LARVAE

The larvae are pelagic, free-floating, and 4-9 mm

long (Das 1972; Graham and Chenoweth 1973; Cooper et

al. 1975). The larval stage of fall-spawned herring in the

Gulf of Maine lasts 4-8 months, depending on the timing

of spawning. The larval stage is shortest for earlyspawned (August) larvae, and longest for late-spawned

(December) herring. Currents affect the pelagic larvae;

however, they may or may not disperse randomly from

the spawning grounds. Some larvae are retained for

several months after hatching on or near the spawning

site, while other larvae are dispersed soon after hatching

and drift with residual currents (Iles and Sinclair 1982;

Townsend et al. 1986; Chenowith et al. 1989; Smith and

Morse 1993).

Larvae from Nantucket Shoals and Georges Bank

tend to drift to the southwest (Lough et al. 1980; Grimm

1983). Larvae produced off southwestern Nova Scotia

are retained initially near the spawning ground and then

drift up into the Bay of Fundy (Iles 1971; Stephenson and

Power 1988). Larvae produced in coastal Gulf of Maine

generally remain inshore (Graham 1982; Townsend 1992)

and disperse in a westerly direction and enter bays and

estuaries where they overwinter (Graham et al. 1972;

Chenoweth et al. 1989; Townsend 1992). In some years,

late-hatched larvae from Jeffreys Ledge and Stellwagen

Bank are transported eastward and overwinter in the

Sheepscot River (Lazzari and Stevenson 1992). During

the first winter after hatching, herring larvae are exposed

to extremely low temperatures and food levels (Townsend

and Graham 1981; Graham et al. 1990). It is not clear if

larval survival is enhanced as a result of overwintering in

nearshore and estuarine waters (Graham 1982) or in

coastal waters (Townsend 1992).

Herring are one of the few species that perform

extensive vertical migrations as larvae. They make diel or

semi-diel vertical migrations throughout the water column

that may be linked to time of day or turbidity (related to

light level), tidal currents, or shifts in prey abundance

(Lough and Cohen 1982). Vertical movements may be a

larval retention mechanism enabling them to control their

displacement by tidal currents (Graham 1972; Stephenson

and Power 1988).

JUVENILES

Larvae metamorphose into juveniles at 40-50 mm

total length (TL) in early spring (April-May). Juveniles

form large schools in coastal waters throughout the Gulf

of Maine (Munroe, in prep.) and off southern New

England, where they have been collected in surveys off

Connecticut and southern Massachusetts in May and June

(A.B. Howe, Massachusetts Division of Marine Fisheries,

East Sandwich, MA, personal communication). In the

summer and fall, juveniles move out of nearshore waters

to overwinter in deep bays or near the bottom in offshore

areas (Boyar 1968). Two-year old juveniles return

inshore the following spring when they are fully recruited

to the coastal fishery.

Juveniles (and adults) perform vertical migrations

that are linked to changing light intensity, most likely in
Page 2

response to movements of their prey (Blaxter 1985).

They move up in the water column at twilight and remain

near the surface when light intensity is low (Johnson

1940; Brawn 1960a); activity is highest just after sunrise

and just before sunset. Blaxter (1985) suggested that

herring move away from the surface in daylight to avoid

predation by diving birds.

ADULTS

Both males and females generally mature between

25-27 cm (O’Brien et al. 1993). Mean lengths of herring

on Georges Bank ranged from 23.7-25.6 cm at age 3 to

33.0-33.3 cm at age 7 (Boyar 1968). Maximum size is

about 39 cm TL and 0.68 kg, and maximum age is 15-18

years (Anthony 1972). Adults almost invariably occur in

large schools. Vertical migrations linked to changing

light intensity are pronounced and are probably related to

movements of prey and avoidance of predatory seabirds

(Blaxter 1985).

A reduction in mean weight at age of adults has

occurred since 1983. The mean weight of fish averaged

across ages 3 to 7 was 247 g in 1983, 160 g in 1988, 137

g in 1994, and 146 g in 1997. Changes in the seasonal

distribution of fishing and changes in the contribution of

faster-growing Georges Bank fish did not affect the

reductions in mean weight because the fishery has

occurred only in the Gulf of Maine since 1983 (D.K.

Stevenson, Maine Deptartment of Marine Resources,

West Boothbay Harbor, ME, personal communication).

REPRODUCTION

In general, males and females mature at around 3-4

years old. Length at maturity of herring has remained

fairly constant for 40 years (Table 1) in contrast with

other New England marine fish species that have

experienced significant declines in size at maturity in

recent years. In this report, size at maturity follows

O’Brien et al. (1993) and lengths were rounded to the

nearest whole centimeter. Thus, herring ≥ 25 cm are

considered adults.

Age at maturity may be density dependent; a higher

percentage of age 3 fish mature when abundance is low

(Tupper et al., in prep.). Beginning in 1983, coincident

with increasing population size (stock recovery), herring

growth rates decreased and the percentage of fish

maturing at age 3 declined, especially on Georges Bank

and Nantucket Shoals (D.K. Stevenson, Maine

Deptartment of Marine Resources, West Boothbay

Harbor, ME, personal communication). The percent of

mature age 3 fish declined from 50-70% in the mid-1980s

to 10-30% in 1990-1996.

Historically, three herring spawning stocks have been

recognized in the U.S. fishery: southwestern Nova Scotia,

coastal Gulf of Maine, and Georges Bank/Nantucket

Shoals (Figure 2). Spawning off Nova Scotia occurs in

the Trinity Ledge/Lurcher Shoals/German Bank area

(Stephenson and Power 1988). In the inshore coastal

areas of the Gulf of Maine, spawning occurs in Scots Bay

in the Bay of Fundy, off eastern Maine and the southwest

shore of Grand Manan Island, off Penobscot Bay, and in

the western gulf off Wood Island, Jeffreys Ledge, and

Stellwagen Bank (Tupper et al., in prep.). On Georges

Bank, major spawning sites have historically been located

near the Northeast Peak, Cultivator Shoals, and Nantucket

Shoals (Boyar 1968; Anthony and Waring 1980; Grimm

1983; Lough et al. 1985) (Figure 3).

Gulf of Maine herring spawn in the fall, typically

between July and November (Sinclair and Tremblay

1984). Spawning begins in the northern areas of the Gulf

and occurs progressively later with decreasing latitude;

spawning commences last on Nantucket Shoals (Bigelow

and Schroeder 1953). Spawning off southwestern Nova

Scotia occurs from July to November and peaks in

September-October (Boyar 1968; Das 1968, 1972). In the

coastal Gulf of Maine, spawning occurs from August to

October (Kelly and Stevenson 1985), and peaks in midSeptember to mid-October in eastern Maine and in

October in western Maine (Graham et al. 1972). On

Jeffreys Ledge, spawning occurs from September to

November (Kelly and Stevenson 1985). On Georges

Bank, spawning occurs from late August to December

(Boyar 1968; Berenbeim and Sigaev 1978; Lough et al.

1980) with a peak in September-October (Boyar 1968;

Pankratov and Sigaev 1973; Grimm 1983). On Nantucket

Shoals, spawning peaks from October to early November,

1-2 weeks later than on Georges Bank (Lough et al. 1980;

Grimm 1983).

There is some evidence of spring spawning.

Approximately 2% of the fish sampled in the coastal Gulf

of Maine and on the southwestern Scotian Shelf during

spring were in spawning condition (Boyar 1968).

FOOD HABITS

Larvae begin exogenous feeding before the yolk sac

is completely absorbed (Bigelow and Schroeder 1953).

They feed opportunistically on whatever zooplankton of

appropriate size are abundant (Sherman and Perkins

1971). Their primary prey are copepods (Bigelow and

Schroeder 1953; Sherman and Honey 1971), in particular,

Pseudocalanus sp., Paracalanus parvus, and Centropages

typicus (Cohen and Lough 1983). Juveniles feed on up to

15 different groups of zooplankton; the most common are

copepods, decapod larvae, cirriped larvae, cladocerans,

and pelecypod larvae (Sherman and Perkins 1971).

Adults have a diet dominated by euphausiids,

chaetognaths, and copepods (Bigelow and Schroeder

1953). Maurer (1976) reported that the most important

prey items of adult herring collected on Georges Bank
Page 3

were chaetognaths (Sagitta elegans, 43% by weight),

euphausiids (Meganyctiphanes norvegica, 23%;

Thysanoessa inermis, 6.1%), pteropods (Limacina

retroversa, 6.2%), and copepods (3%).

During 1973-1980, the diets of juveniles and adults

collected in the Gulf of Maine during Northeast Fisheries

Science Center (NEFSC) trawl surveys were dominated

by euphausiids (47% by weight), of which more than 50%

were M. norvegica, and copepods (26%) (Figure 4a). On

the Scotian Shelf, euphausiids composed more than 50%

of the herring diet. During 1981-1990, amphipods were

the most common prey item on Georges Bank, followed

by mysids (Figure 4b). Present in smaller amounts were

euphausiids, copepods, chaetognaths, and unidentified

fish larvae. Herring diets in southern New England and

the Mid-Atlantic during 1981-1990 were more varied.

PREDATION

Juvenile and adult herring are preyed on by many

marine species, including sand lance (Ammodytes

hexapterus), cod (Gadus morhua), pollock (Pollachius

virens), haddock (Melanogrammus aeglefinus), silver

hake, white hake (Urophycis tenuis), striped bass,

mackerel, billfish, tuna, salmon, sculpins, winter flounder

(Pseudopleuronectes americanus), dogfish, porbeagle

shark, and skates (Raja spp.). Fish predation can be a

significant source of mortality, especially at spawning

time. Several fish species, as well as American lobster

(Homarus americanus) and starfish, eat herring eggs.

Sand lance may consume large quantities of eggs and

larvae, which are sometimes cannibalized by adult herring

as well. Jellyfish may also be an important predator on

the early life stages. Large numbers of herring are also

eaten by marine birds, northern shortfin squid, seals,

porpoises, and whales (Munroe, in prep.).

MIGRATION

Adult herring make extensive feeding, spawning, and

overwintering migrations. Schooling behavior begins at

metamorphosis (Sindermann 1979). Schools are usually

composed of fish of similar size (Bigelow and Schroeder

1953), and to a large extent, of the same year class

(Munroe, in prep.). In the Gulf of Maine, juveniles spend

the summer in inshore areas off Maine and New

Brunswick. In autumn, they move south to waters off

Massachusetts and Rhode Island; they return to Maine the

following spring (Tupper et al., in prep.). Some juveniles

spend at least the spring and early summer off southern

New England, especially off southern Massachusetts

(through at least mid-June) before moving into the Gulf of

Maine or offshore, presumably east of Cape Cod.

Juveniles are sometimes abundant in winter and spring in

the Hudson-Raritan estuary and in fall in Long Island

Sound. Young-of-the-year herring are not effectively

retained by standard resource survey trawls, but in Long

Island Sound, 15-min tows using a trawl with 0.25-inch

codend liner have yielded up to 80,000 herring

(Gottschall et al., in review).

Adult herring are highly migratory and there is

evidence of intermixing of adults from different spawning

groups during the non-spawning phase of their seasonal

cycle (Sinclair and Iles 1985). Three general migratory

patterns are recognized off the northeast coast of the U.S.

(NAFO regions 4X, 5, and 6) (Sindermann 1979; Figure

5). Herring that spend the summer and fall in southwest

Nova Scotia overwinter in Chedabucto Bay in northeast

Nova Scotia. The Georges Bank/Nantucket Shoals stock

overwinters south of Cape Cod and along the midAtlantic coast. The stock moves north onto Georges Bank

and into the Gulf of Maine in the spring before

congregating on spawning grounds southeast of

Nantucket and on Georges Bank in the fall. The

migrations of coastal adults are less well known. Adults

in the western Gulf of Maine may migrate southwest

along the coast after spawning and overwinter at the

western extreme of their migratory path, possibly south of

Cape Cod. Adults in the eastern Gulf of Maine may

migrate southwest and overwinter in Massachusetts Bay

and southern New England.

STOCK STRUCTURE

Atlantic herring may have the most complex stock

structure of any marine fish (Iles and Sinclair 1982) and

attempts to define stock structure have a long history

(Kornfield et al. 1982). Herring in the Gulf of Maine

region have historically been considered three distinct

spawning stocks: Nova Scotia, coastal Gulf of Maine, and

Georges Bank/Nantucket Shoals (Iles 1972; Atlantic

States Marine Fisheries Commission 1995) (Figure 2). In

U.S. waters, they are treated as one coastal stock complex

for assessment (Northeast Fisheries Science Center 1996).

Evidence for and against the discreteness of local herring

stocks includes spawning and larval distributions, tagging

studies, morphometrics and meristics, genetics, and

parasites.

Genetic studies indicate that herring spawning groups

are not discrete, genetically distinct stocks. Safford and

Booke (1992) did not find consistent differences between

herring from two well-separated spawning areas, Jeffreys

Ledge and Trinity Ledge, using traditional enzyme

electrophoresis. Analysis of mitochondrial DNA also

failed to distinguish between fish from these areas

(Kornfield and Bogdanowicz 1987). Kornfield et al.

(1982) found low levels of genetic heterogeneity among

fall spawning herring in the Gulf of St. Lawrence and the

Gulf of Maine, and concluded there is only one genetic

population of fall spawners in the northwest Atlantic.

They did, however, find that spring spawning herring
Page 4

from the Gulf of St. Lawrence were genetically distinct

from fall spawners in the Gulf of St. Lawrence and the

Gulf of Maine.

There has been speculation that adult herring return

to spawn at the spawning grounds where they were born,

but this has only been demonstrated in one study. Herring

off Newfoundland were shown to have a homing rate of

66-93% (Wheeler and Winters 1984). The inability to tag

herring larvae has made it impossible to determine

whether individuals are actually returning to the site

where they were spawned. Results from an international

herring tagging program and from the Canadian

Department of Fisheries and Oceans (summarized in

Stobo 1983) indicate that stocks are generally mixed

throughout most of the year and that spatial and temporal

isolation occurs chiefly during spawning. However,

migration patterns of individual stocks persist among

years and there is little straying of fish from a given stock.

Tagging along the Maine and New Hampshire coasts by

the Maine Department of Marine Resources showed

consistency of migration patterns over time (Creaser and

Libby 1988). Patterns were similar for juveniles and

adults, but adults often covered greater distances; many

adults tagged in summer in eastern Maine overwintered in

Massachusetts Bay. There was some tendency for adults

tagged in eastern Maine to be recovered in the

southwestern Nova Scotian fishery.

Herring in the Gulf of Maine and on Georges Bank

and the Scotian Shelf spawn in well-defined areas,

although homing to natal spawning grounds has not been

demonstrated. Distinct and spatially stable larval

retention areas may also promote genetic isolation. Iles

and Sinclair (1982) stated that larval herring were

concentrated in such areas in the northwest Atlantic and

hypothesized that the number of retention areas

determined the number of genetically distinct stocks.

However, Smith and Morse (1993) discussed evidence for

larval drift in the region and questioned whether stocks

could be separated through larval retention. Chenoweth

et al. (1989) reported extensive westward transport of

larvae from Gulf of Maine spawning sites and possible

larval retention by a sharp oceanographic front near

Grand Manan Island.

Pectoral fin ray counts were once considered the

most promising meristic character for discriminating

stocks. The number of pectoral fin rays is related to water

temperatures and is determined at an early age. Adult

herring from Georges Bank-Cape Cod have lower

pectoral fin ray counts than adults from waters to the

north, presumably due to warmer temperatures. In the

1958-1963 year-classes, herring from eastern Maine and

Nova Scotia had the highest pectoral fin ray counts and

fish from western Maine were intermediate in fin ray

numbers. However, juvenile fish from Maine had counts

similar to fish from Georges Bank-Cape Cod, indicating

that they probably came from that area. It is likely that

some of those juveniles subsequently entered the Georges

Bank fishery (Anthony 1981).

Significant phenotypic differences have been

identified among herring spawning groups, but this may

reflect different environmental histories rather than

genetic differentiation. Safford and Booke (1992) found

differences in several morphometric characters between

herring from Jeffreys Ledge and Trinity Ledge, but

overall results supported the single-population hypothesis.

They postulate that either sufficient gene flow exists

between spawning groups to prevent the evolution of

genetically distinct stocks, or that genetic isolation is a

recent phenomenon and genetic differences have not had

time to evolve.

Parasites may be useful as biological indicators to

differentiate between fish populations. Parasites of

Georges Bank and Gulf of Maine herring have apparently

not been studied, but Bay of Fundy, Scotian Shelf, and

Gulf of St. Lawrence fish have been surveyed

(McGladdery and Burt 1985). Seven of 18 parasite

species identified were potential indicators. Patterns of

occurrence of the parasites indicated movement of fish to

and from the Bay of Fundy, and extensive mixing of

stocks in feeding and nursery areas.

McQuinn (1997) reviewed arguments for a discrete

versus dynamic balance population concept for Atlantic

herring. He proposed that the population structure and

dynamics of herring fit well within a metapopulation

model. This model allows for significant mixing and

gene flow among units that still retain considerable

persistence and discreteness due to behaviorally-induced

homing to spawning grounds. Although the

metapopulation (or stock complex) is the practical unit for

management, local populations must be conserved to

preserve spawning potential and viable coastal fisheries.

The metapopulation may increase resilience of local

populations because a strong year class may enhance

several local populations (McQuinn 1997).

HABITAT CHARACTERISTICS

Information on the life history and habitat

characteristics of Altantic herring are presented here and

are summarized in Tables 2-5. This information is limited

to the Georges Bank, coastal Gulf of Maine, and Nova

Scotia stocks, which occur in U.S. waters at some time

during the year. Information for other stocks in the

northwest and northeast Atlantic were not considered.

EGGS

Herring eggs are usually spawned on horizontal beds

at depths of 40-80 m on Georges Bank, 20-50 m in coastal

Gulf of Maine, and as shallow as 11-13 m off southwest

Nova Scotia. Eggs are laid on gravel (the preferred

substrate), sand, rocks, shell fragments, aquatic
Page 5

macrophytes, and structures such as lobster pots.

Spawning occurs in areas of well-mixed water with tidal

currents of 1.5-3.0 knots. These high energy

environments provide aeration and reduce siltation and

accumulation of metabolites. Spawning occurs at

temperatures of 12-15o

C on Georges Bank, 6-13oC on

Nantucket Shoals, and 8-12o

C near Grand Manan Island,

and at salinities of 31.9-33.0 ppt. Laboratory studies

found normal egg development and hatching at 10 and

15o

C, no development at 0 and 5o, and rapid initial

development followed by 100% mortality at ≥ 20o

C.

LARVAE

Larvae occur at temperatures of 9-16oC and salinities

of 32 ppt in the Gulf of Maine. Survival and growth in

winter may be enhanced in offshore waters, which are up

to 5o

C warmer than inshore waters. Larvae may

acclimate to lower temperatures when the rate of

temperature decline is slow; in the laboratory, survival

was ≤ 30% when the rate of change was 0.1-0.25o

C/day,

but up to 70% when the rate of change was < 0.1o

/day.

Larvae occur at depths > 50 m on Georges Bank where

they are retained in the clockwise current gyre for several

months. Light, turbidity, and tidal currents may control

their vertical migrations.

In the NEFSC Marine Resources Monitoring,

Assessment and Prediction (MARMAP) survey, most

larvae were collected at 8-14o

C from September to

November; maximum abundance was at 9-12o

C (Figure

6). In December, larvae occurred at 6-11o

C with the

majority collected at 8-9o

C. Temperatures at the time of

collection decreased each month from January to March

and increased from April to August. Larvae were

collected at stations with bottom depths ranging from 10-

250 m, although most were collected at stations with

depths of 50-90 m (Figure 6).

JUVENILES

In the Sheepscot River, juveniles prefer temperatures

of 10-16o

C. They may overwinter in Passamaquoddy Bay

until the temperature drops to 0o

C. In the laboratory, the

upper lethal temperature is 19.5-21.2o

C, the lower lethal

temperature is -1.1o

C, and the preferred temperature is 8-

12o

C. Juveniles in the Gulf of Maine occur at average

surface salinities of 31-32.4 ppt. In the Sheepscot River,

they occur at 16-32 ppt, although most occur at 30-32 ppt.

Laboratory studies indicate a general preference for 26-32

ppt. This salinity preference is temperature dependent;

there is a preference for > 29 ppt at < 10o

C. There is a

tendency to prefer higher salinities and to avoid brackish

conditions with increasing fish age.

Juveniles caught during NEFSC bottom trawl surveys

were most abundant at temperatures of 3-4o

C and depths

of 30-90 m in spring, 6-9o

C and 15-135 m in summer, 8oC

and 30-60 m in fall, and 2-4o

C and 30-60 m in winter

(Figure 7a). There is a bimodal distribution in occurrence

relative to temperature based on Massachusetts inshore

trawl survey catches; relative abundance was highest at 4-

7o

C and 12oC in spring, and 7-12oC and 17oC in the fall

(Figure 8). The bimodality may be related to temperature

differences north relative to south of Cape Cod. Relative

abundance was greatest at bottom depths of 5-30 m in

spring and 5-75 m in the fall. In Narragansett Bay,

juveniles were most abundant at 3-6o

C in winter, 10-12oC

in spring, 17-19o

C in summer, and 18-20oC in fall (Figure

9a). Relative abundance was high at bottom depths of

100 ft (30 m) in all seasons and at 30 and 60 ft (9 and 18

m) in spring. In the Hudson-Raritan Estuary, herring

were found at 2-6o

C and 12-22oC but were most abundant

at 4-6o

C and at 15-18oC (Figure 10a). There were few

differences in abundance over the range of depths and

salinities sampled.

ADULTS

In the Gulf of Maine, herring spawn at 7-15oC.

Spawning begins earlier in years when August water

temperatures are warmer. Adults may overwinter at

temperatures as low as 0o

C in Passamaquoddy Bay. They

generally occur at salinities > 28 ppt and spawn at 31.9-

33.0 ppt (never in brackish water). The distribution of

schools is often related to concentrations of their

euphausiid prey; areas with phytoplankton blooms may be

avoided.

Catches of adult herring in the NEFSC bottom trawl

survey were greatest at 5o

C and 30-50 m in spring, 6oC

and 20-130 m in summer, 5-6o

C and 60-170 m in fall, and

7-8o

C and 70-100 m in winter (Figure 7b). In the

Massachusetts inshore trawl survey, the largest catches

occurred at 4-6o

C in depths of 5-75 m in spring and at 7oC

in depths of 50-80 m in the fall (Figure 8). Abundance in

Narragansett Bay was highest at 3-6o

C and 100 ft (30 m)

in winter, 3-5o

C and 100 ft in spring and 7-11oC, and 30 ft

(9 m) in fall; no adults were caught in summer (Figure

9b). In Long Island Sound, springtime abundance was

highest at 9-10o

C, 10-30 m, and salinities of 25-28 ppt.

The largest autumn catches occurred at 17-21o

C, 10-18 m,

and 27-28 ppt. In the Hudson-Raritan estuary, catches

were highest at 3-6o

C and 15-45 ft (4.5-13.5 m) (Figure

10b).

GEOGRAPHICAL DISTRIBUTION

EGGS

The eggs of herring are demersal and adhere to the

substrate (Bigelow and Schroeder 1953; Fahay 1983) and

were not usually collected during the NEFSC MARMAP
Page 6

survey. The general location of herring spawning areas in

the northwest Atlantic Ocean is presented in Figure 2.

LARVAE

The NEFSC MARMAP survey collected herring

larvae from New Jersey to the Bay of Fundy inshore to

the seaward limit of the survey (Figure 11). Larvae were

collected in all months, even though herring in the Gulf of

Maine do not spawn in the spring and larvae undergo

metamorphosis in April and May. The highest mean

monthly density (351 larvae/10 m2

) occurred in

September off southwestern Nova Scotia (Figure 11)

when larvae were restricted to the northeastern Gulf of

Maine. Larvae were relatively abundant in October (39

larvae/10 m2

) and November (49 larvae/10 m2); high

larval densities occurred from the western Gulf of Maine

and Massachusetts Bay to western Georges Bank and

Nantucket Shoals indicating that spawning began earlier

in the northeast (see also Bigelow and Schroeder 1953;

Tupper et al., in prep.). Mean densities were much lower

(less than 6 larvae/10m2

) from December through August.

Herring spawn in the fall (Sinclair and Tremblay 1984)

and with a peak from September to October (Boyar 1968).

The distribution of herring larvae changed

considerably around Georges Bank from 1971 to 1990, a

period of widely fluctuating adult spawning biomass

(Figure 12; Smith and Morse 1993). In 1971, herring

spawned throughout Georges Bank and Nantucket Shoals;

the principal spawning ground was on the Northeast Peak

of Georges Bank. Following the collapse of the Georges

Bank fishery, spawning was restricted to Nantucket

Shoals by 1976. By 1979, larvae were found only around

Stellwagen Bank in Massachusetts Bay. The

reappearance of larvae on Nantucket Shoals in 1985

indicates an increase in spawning stock distribution. By

1988, larvae were collected on Cultivator Shoals on

Georges Bank, but were not found on the Northeast Peak

through 1990.

JUVENILES AND ADULTS

NEFSC Bottom Trawl Survey

The seasonal distribution pattern and abundance of

juvenile and adult herring were similar. Juveniles and

adults range from south of Cape Hatteras to the Bay of

Fundy and Browns Bank (Figure 13). In spring, juveniles

and adults were most abundant on the inner shelf from

North Carolina to New Jersey, shelf-wide from Long

Island to Cape Cod, and in Massachusetts Bay, and

moderately abundant on Georges Bank. Juveniles were

also abundant along the coast of Maine. In summer,

juveniles and adults occurred most frequently in the Gulf

of Maine and to a lesser extent on Georges Bank.

Densities in autumn were highest in Massachusetts Bay,

on northern Georges Bank, and Nantucket Shoals. In

winter, herring were caught throughout the Middle

Atlantic Bight and on southern Georges Bank; juveniles

also occurred in the Gulf of Maine. These distributions

show the overwintering migrations to areas south of Cape

Cod (Tupper et al., in prep.).

Massachusetts Inshore Trawl Survey

In spring, juvenile herring were most abundant

northwest of Cape Ann, throughout Cape Cod Bay, along

the northern shore of Nantucket Island and southern shore

of Martha’s Vineyard, and in Buzzards Bay (Figure 14).

In the fall, the largest catches occurred around Cape Ann,

in central and western Cape Cod Bay, off Buzzards Bay,

and off the southern shore of Martha’s Vineyard. Adults

were most abundant in northern Cape Cod Bay and

around Cape Ann in spring and fall.

Rhode Island Trawl Survey

Catches of juveniles were patchy in Narragansett Bay

(Figure 15). Catches were highest in summer when the

largest mean catch (254 fish/tow) occurred at the station

farthest offshore and five of the 12 stations in the bay had

> 100 per tow. Abundance was lower during the

remaining seasons. Adults were scarce in winter when the

highest mean catch was 12 per tow. Catches were smaller

in other months and no adults were caught in summer.

Connecticut Fisheries Division Survey

In spring, herring were abundant in central Long

Island Sound (Figure 16). Juveniles were not separated

from adults, but most fish were 26-30 cm long (i.e.,

adults). Catches were much smaller in autumn and

occurred mostly along the west-central coast. Most fish

in autumn were 9-12 cm (Gottschall et al., in review).

Hudson-Raritan Estuary Trawl Survey

Catches of all sizes of herring were distributed fairly

evenly throughout the Hudson-Raritan estuary (Figure

17). Juveniles were most abundant in winter and spring

throughout the lower estuary. They were sometimes

common at the mouth of the estuary in summer, and were

rare in fall. Adults were most common in winter, which

is consistent with the fact that adults from the Gulf of

Maine overwinter south of Cape Cod (Sindermann 1979;

Tupper et al. in prep.). Adult herring were occasionally

collected throughout the survey area in spring and fall,

however none were caught in summer.
Page 7

Estuarine Living Marine Resources

The NOAA Estuarine Living Marine Resources

Program (ELMR) compiled information on the

distribution and abundance of all life stages of Atlantic

herring in estuaries in the New England (Jury et al. 1994)

and the Middle Atlantic (Stone et al. 1994) (Table 6).

Adults and juveniles were ‘highly abundant’ in the

northernmost estuaries (Passamaquoddy Bay through

Penobscot Bay). Larvae were ‘highly abundant’ from

Englishman-Machias Bays through the Sheepscot River.

Abundance of all life stages was lower in the Middle

Atlantic estuaries; only adults were abundant in

Narragansett Bay (Jan-Apr), Long Island Sound (NovMay) and Great South Bay, Long Island (Nov-Feb).

Herring occurred in all major estuaries south to

Chesapeake Bay.

STATUS OF THE STOCKS

Atlantic herring were extremely abundant in

northeastern U.S. waters during the 1960s and were fished

intensively by a large foreign fleet. The Georges BankNantucket Shoals fishery extracted a peak of 373,598 mt

in 1968, and an average of 168,750 mt/year over a 16-

year period before the stock collapsed in the early 1970s

(Atlantic States Marine Fisheries Commission 1995).

Landings remained low for about 10 years, but stock

biomass is now high and apparently increasing (Figure

18; Northeast Fisheries Science Center 1996). The stock

complex is under-utilized (Northeast Fisheries Science

Center 1996), but the Gulf of Maine portion of the

complex may be fully exploited (Atlantic States Marine

Fisheries Commission, unpublished data).

RESEARCH NEEDS

Historically, Atlantic herring in the Gulf of Maine

have supported large, economically important fisheries

(Friedland 1995). Herring have a complex life history

and many areas still require study. The Gulf of Maine

Aquarium Development Corporation has identified

several research needs for Gulf of Maine herring (Tupper

et al., in prep.):

• Identify discrete populations/metapopulations and

major and minor spawning components in the Gulf of

Maine/Bay of Fundy region and the degree of

intermixing. Consider using scales, otolith structure,

and possibly morphometrics. Concentrate on

spawning grounds and tag ripe and running fish only.

Perhaps combine with acoustic surveys.

• Explore new technologies (e.g., acoustics or laser

illumination) for improving surveys of all life stages.

• Validate the current natural mortality estimate for

Gulf of Maine/Bay of Fundy (18%). Synthesize

information on mammal, seabird, and other

predation. Examine size/age-specific natural

mortality. Identify oceanographic influences on

larval survival, particularly effects of temperature,

climate change, and plankton patch dynamics.

• Conduct surveys to provide an overview of larval

abundance/distribution throughout the Gulf of Maine

for a single year. Determining the fate of herring

spawned on Jeffreys Ledge is a high priority.

Other research needs that became apparent during

development of this report include:

• Sample the Northeast Peak of Georges Bank for

larvae, which were present there 30 years ago but not

in 1990.

• Conduct experimental studies of temperature and

salinity preferences; most existing information is for

European stocks.

• Prior attempts to discriminate stocks by analyzing

otolith elemental composition have been

unsuccessful, but given recent improvements in

analytical techniques this line of research may now

be more promising.

• Map the distribution of seabed habitat types,

including determining the scale of detail needed for

habitat mapping.

• Continue efforts to locate all significant herring

spawning areas.

• Determine effects of bottom-tending fishing gears

and natural processes on spawning grounds.

• Determine the value of marine protected areas for

conserving and enhancing herring stocks. Identify

how these areas would function as larval exporters

and collectors.

ACKNOWLEDGMENTS

Bruce Collette provided the draft update to the

Bigelow and Schroeder (1953) section on herring written

by Thomas Munroe. Judy Berrien, Rande Ramsey-Cross,

and Claire Steimle aided with the literature search.

Sukwoo Chang assisted in generating the maps and

histograms from NEFSC trawl surveys. Stuart Wilk,

Robert Pikanowski, and Donald McMillan produced the

maps and histograms for the Hudson-Raritan data and

Jeffrey Cross produced the status of the stocks figure.

Helpful comments on drafts of the document were

provided by David Stevenson, Kevin Friedland and

technical experts associated with the New England

Fishery Management Council.

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Page 12

Table 1. Size and age at sexual maturity for Atlantic herring, Clupea harengus.

Period Age at Maturity

(A50, years)

Size at Maturity

(L50, cm)

Reference

male female male female

1987-1989 2.9 3.0 25.3 25.4 O’Brien et al. (1993)

1966-1975 - - 25.4-27.4 Sinclair et al. (1982)

1949-1952 - - 26.9 Scattergood (1952)
Page 13

Table 2. Summary of life history and habitat parameters for Atlantic herring, Clupea harengus – Georges Bank.

Life Stage Size and Growth Habitat Substrate Temperature Salinity

Eggs 1

Spawning beds level

and horizontal with

only occasional

shallow depressions or

ridges; at 40-80 m

depth. Major

spawning site on NE

part of Georges Bank.

Herring spawn found

only on gravel (2-10

mm diameter)

associated with strong

bottom currents.

Although gravel

substrate is extensive,

egg beds are limited to

small region on

western edge of

northeast area of the

Bank.

Temperatures in the

vicinity of the Georges

Bank egg beds tend to

be 12-15o

C. Nantucket

Shoals tends to be

colder (6-13o

C). There

was an increase of 2-

3o

C on Georges/Gulf

of Maine from the late

1960s to 1977.

High salinities of 32 ppt

reported in surface

waters around egg beds.

Larvae 2

Recently hatched:

4-9 mm TL, mean = 7

mm.

Total size range:

4-45 mm.

Growth = 0.2 mm/4

days.

Adults 3

(age: avg. length, cm)

III: 23.7 - 25.6

IV: 27.1 - 27.9

V: 28.9 - 29.4

VI: 30.6 - 30.8

VII: 31.4 - 32.1

VIII: 33.0 - 33.3

Spawn on gravel sea

floors; attachment of

eggs to stable material

prevents translocation

by strong currents.

Correlation has been

demonstrated between

summer thermal

regime (i.e.,

temperature in August)

and the date of peak

spawning; a warm

August results in an

earlier spawning peak.

1 Boyar (1968), Caddy and Iles (1973), Drapeau (1973), Graham and Chenoweth (1973), Pankratov and Sigaev (1973), Berenbeim and Sigaev (1978),

Lough et al. (1980, 1985), Grimm (1983), Valentine and Lough (1991) 2

 Boyar et al. (1973), Graham and Chenoweth (1973), Lough et al. (1980, 1985), Cohen and Lough (1983), Grimm (1983) 3

 Boyar (1968), Drapeau (1973), Pankratov and Sigaev (1973), Maurer (1976), Berenbeim and Sigaev (1978)
Page 14

Table 2. cont’d.

Life Stage Currents Prey Predators Notes

Eggs 1

High energy environments;

tidal action provides aeration,

prevents siltation and

accumulation of metabolites.

N/A

Increased abundance of other

fish species in areas of

spawn; 4 most common fish:

red hake, sculpin, dogfish,

skate; also increase in

starfish and moon snails.

1-2 cm (7-14 layers) thick

egg mat. Area of egg bed

ranges from 4500 to 10000

km2

. Egg mortality varies:

on north and south spawning

beds, approx. 8% of spawn

removed within 1-2 days of

hatching. Surveys on eastern

Georges Bank over 5

spawning seasons (1964-

1970) show year to year

decrease in area occupied by

egg beds.

Spawning time: late Aug -

Oct; peaks in mid/late Sept -

Oct.

Larvae 2

Clockwise current gyre;

larvae generally dispersed in

a SW direction (2-15 km/d),

towards coastal Gulf of

Maine.

Primary prey: juvenile stages

and adults of seasonally

dominant copepods.

The 3 most important species

were Pseudocalanus sp.,

Paracalanus parvus and

Centropages typicus. Feeding

activity peaked twice daily:

shortly after sunrise and in

mid-afternoon.

Northeast Georges Bank:

highest larval abundance on

Bank; maximum abundance

mid-late Oct.

Nantucket Shoals: maximum

larval abundance late Oct -

early Nov.

> 80% larval production

occurred on Nantucket

Shoals in 1976-1978.

Estimated larval mortality in

NW in 5 day period = 75%

(< 10 mm); winter mortality

on Maine coast much lower.

Adults 3

High energy environments;

tidal currents and storm

waves.

Primary prey: the

chaetognath Sagitta elegans

(43% by weight); the

euphausiids

Meganyctiphanes norvegica

(23.1%) and Thysanoessa

inermis (6.1%), the pteropod

Limacina retroversa (6.2%),

copepods (3%). May avoid

feeding in areas with

phytoplankton blooms.

Mean size of spawning fish =

29.5 cm; sex ratio 1:1 (Aug -

Sept).

1 Boyar (1968), Caddy and Iles (1973), Drapeau (1973), Graham and Chenoweth (1973), Pankratov and Sigaev (1973), Berenbeim and Sigaev (1978),

Lough et al. (1980, 1985), Grimm (1983), Valentine and Lough (1991) 2

 Boyar et al. (1973), Graham and Chenoweth (1973), Lough et al. (1980, 1985), Cohen and Lough (1983), Grimm (1983) 3

 Boyar (1968), Drapeau (1973), Pankratov and Sigaev (1973), Maurer (1976), Berenbeim and Sigaev (1978)
Page 15

Table 3. Summary of life history and habitat parameters for Atlantic herring, Clupea harengus – Coastal Gulf of Maine.

Life Stage Size and Growth Habitat Substrate Temperature Salinity

Eggs 1

Depths of egg beds varies

from 20 to 50 m,

averaging 45 m; deeper

beds have been shown to

be more successful: 35%

greater egg density and

2X higher mean egg

abundance.

Preferred substrate is

gravel associated

with strong bottom

currents. Gravel

often mixed with

shell fragments

and/or sand; can be

almost 100% shell

fragments.

On Jeffreys Ledge,

eggs spawned on

boulders & rocks,

gravel & coarse sand,

and on red alga

Ptilota serrata .

Hatching success

temperature dependent.

Lab results: 10o

C and 15oC:

egg development &

hatching normal; 0 and

5o

C: no development; 20,

25 and 30oC: rapid

development began but

100% mortality.

Field results: 9.6o

C,

average bottom

temperature at spawning .

Larvae 2

Newly hatched:

4-6 mm

autumn:

7-10 mm

winter:

21-30 mm

spring:

31-40 mm

Larvae may be able to

acclimate to lower winter

temps. when T declines

more slowly. Survival <

30% when T changes 0.1-

0.25o

C/d; more variable

(20-70%) when < 0.1o

C/d.

Low temp. effects may be

avoided through

acclimation & occupancy

of warmer coastal water.

Larvae that

overwinter in

estuaries typically

experience reduced

salinities.

Juveniles 3

4-23 cm Tendency to move to

surface at night results in

increased vulnerability to

fixed gear fishery during

dark phases of the moon.

One study has shown that

juveniles overwinter with

adults in Passamaquoddy

Bay; they remain in the

bay until temp. reaches

0o

C.

Lab: upper lethal = 19.5-

21.2o

C, lower = -1.1oC,

preferred = 8-12 o

C. Field:

preference for 10-16oC in

Sheepscot River; Sardine

production positively

correlated to stock size &

temp., but density overrides

temp. when abundance

high. Highest catches in

the nearshore weir/stop

seine fisheries at 10-13o

C;

> 13o

C activity declines.

Juvenile schools disappear

in colder months (Nov.-

Mar.). Effects of temp. on

determination of yr-class

strength occurs during late

larval/early juvenile phase.

Lab: preference for

26-32 ppt, can resist

salinities as low as 5

ppt for brief periods;

at < 10o

C a

preference for > 29

ppt; at > 10o

C no

salinity preference

seen. Field: present

in 16-32 ppt; highest

abundance at 30-32

ppt. Older juveniles

generally avoid

brackish conditions.

Adults 4

(age: avg. length,

cm) III: 23 - 26

IV: 27 - 28

V: 29 - 30

VI: 30 - 31

VII: 31.9 - 32

VIII: 33 - 33.4

One study has shown that

adults overwinter (along

with juveniles) in

Passamaquoddy Bay;

remain there down to 0o

C.

Spawning in Grand Manan

and northern Gulf primarily

at 8-12o

C.

1 MacFarland (1931),Boyar (1968), Graham et al. (1972), Cooper et al. (1975), Kelly and Stevenson (1985), Townsend et al. (1986), Stevenson and

Knowles (1988), Chenoweth et al. (1989), Stevenson (1989) 2 Sherman and Honey (1971), Graham (1972), Boyar et al. (1973), Cooper et al. (1975), Graham and Townsend (1985), Chenoweth et al. (1989),

Graham et al. (1990) 3

 Brawn (1960a, b, c), Anthony (1971), Stickney (1969), Sherman and Perkins (1971), Recksiek and McCleave (1973), Sindermann (1979), Anthony

and Fogarty (1985)

4 Bigelow and Schroeder (1953),Boyar (1968), Sherman and Perkins (1971), Cooper et al. (1975), Kelly and Stevenson (1985), Munroe (in prep.)
Page 16

Table 3. cont’d.

Life Stage Currents Prey Predators Notes

Eggs 1

Bottom currents at spawning

beds 0 - 1.0 knots.

N/A Level of egg predation varies.

Most abundant predators on

eggs:

1) cunner, Tautogolabrus

adspersus,

2) cod, Gadus morhua.

1-3 cm thick egg mat (20-

30 eggs deep) and low egg

mortality (< 5%) reported.

Egg beds elliptical to

irregular in shape; 2/3 to 1

1/3 km2

in area. 90% of eggs on

rock-gravel. Not known if

Jeffreys herring spawn

selectively over algal

clumps or if algae function

as egg traps. Hatching

success (excluding

predation) 99%. Spawning

time: mid Aug - Nov;

peaks Sept - Oct.

Larvae 2

Use tidal flows to migrate.

On ebb, majority of larvae

shallow; on flood, majority

deep. More larvae at

landward end of channel than

seaward.

Seasonal differences in diet;

prey principally on 5 groups

of zooplankton:

copepods, crustacean eggs,

crustacean nauplii, cirriped

larvae, and tintinnids. Prey

volume (cc/10m3

): summer =

1.1, autumn = 0.5, winter =

0.2, spring = 0.8.

Low temp. may indirectly

increase starvation &

vulnerability to predation.

Selective tidal transport

(larvae retained within

estuary despite seaward

flow). Mortality avg.

2%/d; growth 0.199 mm/d.

No growth difference in

early vs. late spawned

cohorts. Mortality & G

inversely correlated.

Larvae drift from eastern

Maine spawning ground to

estuaries.

Juveniles 3

Opportunistic feeders. 15

groups of zooplankton eaten;

only 5 by > 20% of fish: (1)

copepods, (2) decapod larvae,

(3) cirriped larvae, (4)

cladocerans, (5) pelecypod

larvae; copepods are the most

important food item year

round.

Diurnal vertical movements

in response to changing

light intensity.

Regardless of year class,

western Maine herring

grow faster through age 3

than eastern Maine; at the

end of age 2 avg. ~3 cm

longer. Plankton less

abundant, water temp.

lower, salinity greater in

eastern Maine than in

central or western Maine.

Adults 4

Selective, opportunistic

feeders. Primary prey: shrimp

and copepods. Distribution of

schools of large herring likely

related to presence/absence of

euphausiids.

Spawning adults preyed on by

bluefish and pollock; ranging

from 30-65 cm TL. Predation

mostly at night.

Spawning areas from

Jeffreys Ledge to eastern

Maine should be

considered as a single

spawning population.

1 MacFarland (1931),Boyar (1968), Graham et al. (1972), Cooper et al. (1975), Kelly and Stevenson (1985), Townsend et al. (1986), Stevenson and

Knowles (1988), Chenoweth et al. (1989), Stevenson (1989) 2 Sherman and Honey (1971), Graham (1972), Boyar et al. (1973), Cooper et al. (1975), Graham and Townsend (1985), Chenoweth et al. (1989),

Graham et al. (1990) 3

 Brawn (1960a, b, c), Stickney (1969), Anthony (1971), Sherman and Perkins (1971), Recksiek and McCleave (1973), Sindermann (1979), Anthony

and Fogarty (1985)

4 Bigelow and Schroeder (1953),Boyar (1968), Sherman and Perkins (1971), Cooper et al. (1975), Kelly and Stevenson (1985), Munroe (in prep.)
Page 17

Table 4. Summary of life history and habitat parameters for Atlantic herring, Clupea harengus – Nova Scotia.

Life Stage Size and

Growth

Habitat Substrate Temperature Salinity Currents Prey Predators Notes

Eggs 1 Spawning

bed at Black

Point, NS at

depth of 11-

13 m.

Egg beds on

sand and

small stones.

Temperature of

water column in

spawning area

ranged from

9-15 oC.

Tidal

currents at

egg beds

1.5-2.0

knots.

N/A Haddock is

main egg

predator.

3.25 cm thick

egg bed

reported.

Spawning

time: Aug -

Oct;

peaks late Aug

- Sept.

Larvae 2 Recently

hatched: 5-9

mm TL.

Initial growth

rate =

2 mm/week;

late

autumn/winter

months < 1

mm/week;

spring/early

summer = 2.5

mm/week.

Spawning

site in SW

Nova Scotia

in an area of

well mixed

water.

Temperature of

water column in

spawning area

ranged from

9-15 o

C.

Salinity

ranged

very little

from

32.09 -

32.56 ppt.

Semidiel

pattern of

vertical

migration

demonstrated;

possibly linked

to time of day

(light) and/or

tidal currents.

Adults 3 (age: average

length, cm)

III: 23.7

IV: 26.4 - 27.9

V: 28.9 - 29.6

VI: 30.7 - 30.9

VII: 32.0 - 32.1

VIII: 33.0 -

33.4

1

 McKenzie (1964), Boyar (1968), Das (1968, 1972), Stephenson and Power (1988)

2 McKenzie (1964), Das (1972), Stephenson and Power (1988)

3 Boyar (1968)
Page 18

Table 5. Summary of life history and habitat parameters for Atlantic herring, Clupea harengus – No specific location

given in literature.

Life Stage Size and Growth Habitat Substrate Temperature Salinity

Eggs 1

1.0-1.4 mm in

diameter.

All spawning grounds

in high energy

environments, either

nearshore shallows

subject to wave/tidal

flux, or deeper water

with tidal action.

Spawning substrate

varied (stones, gravel);

free of fine sediments

that might prevent

gaseous exchange

between eggs and

environment.

Bottom temp. of

5-15o

C required.

Average incubation

time for autumn

spawned eggs is 10-15

days. Developmental

rate inversely related

to temp.: 40 d at 4-5o

C,

15 d at 6-8o

C, 11 d at

10-12o

C, 6-8 d at 14.4-

16o

C.

Larvae 2

Occur in 9-16oC in the

Gulf of Maine.

Offshore waters in

winter generally have

higher temperatures

than inshore waters (up

to 5o

C difference); may

favor a more rapid

development in

offshore waters,

thereby reducing time

of vulnerability to

predation.

Juveniles 3

Preference for higher

salinities with

increasing age.

Adults 4

Spawn on stable

material: small stones,

gravel.

Movements become

sluggish at less than

4o

C.

Spawning occurs at

temperatures of 7-

15o

C. Spawning in

western Gulf of Maine

occurs at warmer

temperatures than east.

Enter bays and

estuaries, but 28 ppt is

lower limit of

occurrence.

Spawn at high

salinities, ranging from

31.9 - 33.0 ppt; never

brackish water.

1 Bigelow and Schroeder (1953), Haegele and Schweigert (1985), Munroe (in prep.)

2

 Colton and Byron (1977), Munroe (in prep.)

3 Recksiek and McCleave (1973), Munroe (in prep.)

4 Haegele and Schweigert (1985), Munroe (in prep.)
Page 19

Table 5. cont’d.

Life Stage Currents Prey Predators Notes

Eggs 1

Spawning generally occurs

in areas with good tidal

exchange: average 1.5-3

knots tidal current.

N/A Predation by a variety of

bottom predators (Winter

Flounder major egg

predator). Cannibalism by

adult herring occurs.

Eggs are demersal, adhesive.

Eggs laid in sheets in

successive layers; rarely

exceeds 2 cm in thickness.

Egg mortality is primarily due

to suffocation (from high egg

densities and siltation) and

predation. Spawning time: Sept

- early Nov.

Larvae 2

Begin exogenous feeding

before yolk sac disappears.

Select the most abundant

prey of a suitable size range;

seasonal differences occur.

Primary prey: copepod eggs,

nauplii, copepods, mollusk

larvae. As larvae grow,

consume larger proportion

of copepods.

Solitary and pelagic;

vulnerable to planktonic

predators: jellyfish,

chaetognaths, larger

copepods, euphausiids and

pelagic fishes.

Larvae exhibit diurnal

migratory behavior. Possible

controlling mechanisms: light

level, turbidity, shifts in prey

location & tidal effects.

Juveniles 3

Selective, opportunistic

feeders; predominantly

copepod diet. In darkness:

stop schooling behavior;

swim in tight paths & feed

only by filtering (unable to

feed by biting). In the light:

can feed by either particle

biting or filtering.

Preyed upon by almost all

pelagic predators, including

fishes, marine birds,

northern shortfin squid, and

marine mammals.

Vertical diurnal movements

occur in all seasons. Juveniles

often active near/at the surface

at night; generally move up

water column at dusk.

Adults 4

Selective, opportunistic

feeders; predominantly

euphausiid diet, also

chaetognaths and copepods.

Preyed upon by almost all

pelagic predators, including

fishes, marine birds,

northern shortfin squid, and

marine mammals. Predation

by fish is intense during

spawning.

1 Bigelow and Schroeder (1953), Haegele and Schweigert (1985), Munroe (in prep.)

2

 Colton and Byron (1977), Munroe (in prep.)

3 Recksiek and McCleave (1973), Munroe (in prep.)

4 Haegele and Schweigert (1985), Munroe (in prep.)
Page 20

Table 6. Relative abundance of eggs, larvae, and juvenile Atlantic herring (Clupea harengus) in New England and MidAtlantic estuaries by salinity zone, based on Estuarine Living Marine Resources (ELMR) data in Jury et al. (1994) and

Stone et al. (1994). Salinity zone: T = tidal fresh, M = mixing zone, S = seawater, • = salinity zone not present. Relative

abundance: H = highly abundant, A = abundant, C = common, R = rare, blank = not present.

Eggs Larvae Juveniles

T M S T M S T M S

Passamaquoddy Bay C A A H

Englishman/Machias Bays C A H C H

Narraguagus Bay A H C H

Blue Hill Bay A H C H

Penobscot Bay H H C H

Muscongus Bay A H A A

Damariscotta River A H C A

Sheepscot River A H C A

Kennebec/Androscoggin Rivers C C C C

Casco Bay R A A C A

Saco Bay C A C A

Wells Harbor • • C A • A H

Great Bay C C C C

Merrimack River • C • C •

Massachusetts Bay •• • • A • • A

Boston Harbor • • R A • C A

Cape Cod Bay • R • C • C A

Waquoit Bay R R R

Buzzards Bay R C C

Narragansett Bay C C C

Long Island Sound R R C C

Connecticut River • • R •

Gardiners Bay •• • R C

Great South Bay, NY •• • C

Hudson River/Raritan Bay C C C C

Barnegat Bay, NJ R R C C

New Jersey Inland Bays R R C C

Delaware Bay R R C C

Delaware Inland Bays •• • R

Chincoteague Bay •• • • • • R

Chesapeake Bay Mainstem R

Chester River •••

Choptank River •••

Patuxent River •••

Potomac River •••

Tangier/Pocomoke Sound • •• •• •

Rappahannock River •••

York River, VA •••

James River, VA •••
Page 21

Table 6. cont’d. Relative abundance of spawning adult and adult Atlantic herring (Clupea harengus) in New England and

Mid-Atlantic estuaries by salinity zone based on Estuarine Living Marine Resources (ELMR) data in Jury et al. (1994) and

Stone et al. (1994). Salinity zone: T = tidal fresh, M = mixing zone, S = seawater, • = salinity zone not present. Relative

abundance: H = highly abundant, A = abundant, C = common, R = rare, blank = not present.

Spawning Adults Adults

T M S T M S

Passamaquoddy Bay A H

Englishman/Machias Bays C C H

Narraguagus Bay C H

Blue Hill Bay C H

Penobscot Bay C H

Muscongus Bay C A

Damariscotta River C A

Sheepscot River C A

Kennebec/Androscoggin Rivers C C

Casco Bay R R

Saco Bay R

Wells Harbor • • R C

Great Bay R C

Merrimack River • R •

Massachusetts Bay •• • • A

Boston Harbor • • C A

Cape Cod Bay • R • C A

Waquoit Bay • • R

Buzzards Bay • • C C

Narragansett Bay C A

Long Island Sound C A

Connecticut River • R •

Gardiners Bay • • R C

Great South Bay, NY • • A

Hudson River/Raritan Bay C C

Barnegat Bay, NJ C C

New Jersey Inland Bays C C

Delaware Bay R C

Delaware Inland Bays • • R

Chincoteague Bay •• • •

Chesapeake Bay Mainstem R C

Chester River • •

Choptank River • •

Patuxent River • •

Potomac River • •

Tangier/Pocomoke Sound • •• •

Rappahannock River • R •

York River, VA • R •

James River, VA • R •
Page 22

Figure 1. The Atlantic herring, Clupea harengus L. (from Goode 1884).
Page 23

Figure 2. Location of Atlantic herring spawning populations within the Gulf of Maine area. Solid black represents

spawning areas, while hatched lines represent areas of herring catch (from Iles 1972).
Page 24

Figure 3. Principal spawning grounds on Georges Bank, 1964-1971 (excluding 1967), with a comparison of egg patch

sizes among years (from Anthony and Waring 1980).
Page 25

Figure 4. Abundance (percent of total prey volume) of the major prey items in the diet of Atlantic herring from the

Scotian Shelf, the Gulf of Maine, Georges Bank, southern New England, and the Middle Atlantic based on NEFSC

bottom trawl survey data on food habits, a) 1973-1980 and b) 1981-1990. Methods for sampling, processing, and

analysis of samples differed between the time periods [see Reid et al. (1999) for details]. The category “animal remains”

refers to unidentifiable animal matter.

Euphausiidae 30.8%

Crustacea 25.0%

Animal Remains 24.6%

Thysanoessa raschi 17.3%

Euphausia sp. 1.7%

All Other Prey 0.6%

Meganyctiphanes norvegica 21.6%

Animal Remains 17.0%

Copepoda 15.6%

Crustacea 14.8%

Thysanoessa raschi 10.7%

Euphausiidae 6.6%

Calanoida 5.6%

Parathemisto sp. 2.1%

Hyperiidae 1.9%

Cirripedia 1.4%

Themisto gaudichaudii 1.3%

All Other Prey 1.3%

Animal Remains 63.7%

Copepoda 30.8%

Polychaeta 2.5%

Crustacea Shrimp 2.5%

Cestoda 0.5%

Amphipoda 31.8%

Mysidacea 22.7%

Animal Remains 22.7% Crustacea 9.1%

Pandalidae 4.5%

Euphausiidae 4.5%

Ammodytes sp. 4.5%

Amphipoda 25.6% Animal Remains 17.9%

Chaetognatha 11.1%

Fish 8.6%

Crustacea 8.4%

Mysidacea 8.4%

Copepoda 7.8%

Euphausiidae 5.0%

Gastropoda 3.1%

Thecosomata 1.8%

Decapoda Shrimp 1.3%

Fish Larvae 1.0%

Amphipoda 17.6% Animal Remains 24.9%

Crustacea 10.5%

Mysidacea 8.4%

Ammodytes sp. 8.4%

Euphausiidae 7.1%

Crustacea Shrimp 5.9%

Decapoda Shrimp 4.6%

Chaetognatha 2.7%

Crangon septemspinosa 2.1%

Fish 1.8%

Copepoda 1.8%

Sagitta sp. 1.3%

All Other Prey 2.8%

b) 1981-1990

a) 1973-1980 Scotian Shelf Gulf of Maine

Gulf of Maine Georges Bank

Southern New England Middle Atlantic Bight
Page 26

Figure 5. Hypothesized seasonal movements of three Atlantic herring spawning stocks inhabiting U.S. waters (modified

from Sindermann 1979).
Page 27

Figure 6. Mean water column temperature and bottom depth at stations where Atlantic herring larvae were collected

(solid bars) and at all stations sampled (open bars) during NEFSC MARMAP ichthyoplankton surveys, 1977-1987.

September

0

10

90

100

Stations

Catch

October

0

10

20

40

50

November

0

10

30

40

50

December

Percent

0

10

20

30

40

50

January

0

10

20

30

40

50

February

Water-Column Temperature (0-200m, C)

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

0

10

20

30

40

50

Atlantic Herring Larvae

March

0

10

20

30

40

Stations

Catch

April

0

10

20

30

40

May

0

10

20

40

60

80

June

Percent

0

10

20

40

50

July

0

10

80

100

August

Water-Column Temperature (0-200m, C)

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28

0

10

80

90

Atlantic Herring Larvae

September

0

10

20

90

100

Stations

Catch

October

0

10

20

50

60

November

0

10

20

30

40

December

Percent

0

10

20

30

January

0

10

20

30

40

50

60

February

Bottom Depth (m), Interval Midpoint

10

30

50

70

90

110

130

150

170

190

210

230

250

270

290

325

375

450

750

1250

1750

>2000

0

10

20

30

40

50

Atlantic Herring Larvae

March

0

10

20

30

40

50

60

Stations

Catch

Atlantic Herring Larvae

April

0

10

20

30

May

Percent

0

10

20

30

40

June

0

10

20

90

100

July

0

10

20

30

90

100

August

Bottom Depth (m), Interval Midpoint

10

30

50

70

90

110

130

150

170

190

210

230

250

270

290

325

375

450

750

1250

1750

>2000

0

10

20

30

90

100
Page 28

Figure 7a. Mean bottom water temperature and depth at stations where juvenile (< 25 cm TL) Atlantic herring were

collected (solid bars) and at all stations sampled (open bars) during NEFSC bottom trawl surveys.

FALL

0

5

10

15

20

25

30

35

40

45

50

55

024681

0

1

2

1

4

1

6

1

8

2

0

2

2

2

4

2

6

BOTTOM TEMPERATURE (C)

PERCENT

STATIONS N=788

CATCHES N=43142

WINTER

0

5

10

15

20

25

30

35

40

45

50

55

024681

0

1

2

1

4

1

6

1

8

2

0

2

2

2

4

2

6

BOTTOM TEMPERATURE (C)

PERCENT

STATIONS N=407

CATCHES N=25461

SPRING

0

5

10

15

20

25

30

35

40

45

50

55

024681

0

1

2

1

4

1

6

1

8

2

0

2

2

2

4

2

6

BOTTOM TEMPERATURE (C)

PERCENT

STATIONS N=2293

CATCHES N=129312

SUMMER

0

5

10

15

20

25

30

35

40

45

50

55

024681

0

1

2

1

4

1

6

1

8

2

0

2

2

2

4

2

6

BOTTOM TEMPERATURE (C)

PERCENT

STATIONS N=207

CATCHES N=7822

WINTER

0

5

10

15

20

25

30

35

40

45

50

55

024681

0

1

2

1

4

1

6

1

8

2

0

2

2

2

4

2

6

BOTTOM TEMPERATURE (C)

PERCENT

STATIONS N=407

CATCHES N=25461

FALL

0

5

10

15

20

25

30

35

40

45

0 3

0

6

0

9

0

1

2

0

1

5

0

1

8

0

2

1

0

2

4

0

2

7

0

3

0

0

3

3

0

3

6

0

DEPTH (m)

PERCENT

STATIONS N=928

CATCHES N=52218

WINTER

0

5

10

15

20

25

30

35

40

45

0 3

0

6

0

9

0

1

2

0

1

5

0

1

8

0

2

1

0

2

4

0

2

7

0

3

0

0

3

3

0

3

6

0

DEPTH (m)

PERCENT

STATIONS N=407

CATCHES N=25461

SPRING

0

5

10

15

20

25

30

35

40

45

0 3

0

6

0

9

0

1

2

0

1

5

0

1

8

0

2

1

0

2

4

0

2

7

0

3

0

0

3

3

0

3

6

0

DEPTH (m)

PERCENT

STATIONS N=2697

CATCHES N=145116

SUMMER

0

5

10

15

20

25

30

35

40

45

0 3

0

6

0

9

0

1

2

0

1

5

0

1

8

0

2

1

0

2

4

0

2

7

0

3

0

0

3

3

0

3

6

0

DEPTH (m)

PERCENT

STATIONS N=220

CATCHES N=8109
Page 29

Figure 7b. Mean bottom water temperature and depth at stations where adult (≥ 25 cm TL) Atlantic herring were

collected (solid bars) and at all stations (open bars) sampled during NEFSC bottom trawl surveys.

FALL

0

5

10

15

20

25

30

35

40

45

50

55

024681

0

1

2

1

4

1

6

1

8

2

0

2

2

2

4

2

6

BOTTOM TEMPERATURE (C)

PERCENT

STATIONS N=902

CATCHES N=25161

WINTER

0

5

10

15

20

25

30

35

40

45

50

55

024681

0

1

2

1

4

1

6

1

8

2

0

2

2

2

4

2

6

BOTTOM TEMPERATURE (C)

PERCENT

STATIONS N=351

CATCHES N=16013

SPRING

0

5

10

15

20

25

30

35

40

45

50

55

024681

0

1

2

1

4

1

6

1

8

2

0

2

2

2

4

2

6

BOTTOM TEMPERATURE (C)

PERCENT

STATIONS N=-1820

CATCHES N=40879

SUMMER

0

5

10

15

20

25

30

35

40

45

50

55

024681

0

1

2

1

4

1

6

1

8

2

0

2

2

2

4

2

6

BOTTOM TEMPERATURE (C)

PERCENT

STATIONS N=263

CATCFHES N=8046

FALL

0

5

10

15

20

25

30

35

40

45

0 3

0

6

0

9

0

1

2

0

1

5

0

1

8

0

2

1

0

2

4

0

2

7

0

3

0

0

3

3

0

3

6

0

DEPTH (m)

PERCENT

STATIONS N=1040

CATCHES N=30612

WINTER

0

5

10

15

20

25

30

35

40

45

0 3

0

6

0

9

0

1

2

0

1

5

0

1

8

0

2

1

0

2

4

0

2

7

0

3

0

0

3

3

0

3

6

0

DEPTH (m)

PERCENT

STATIONS N=351

CATCHES N=16013

SPRING

0

5

10

15

20

25

30

35

40

45

0 3

0

6

0

9

0

1

2

0

1

5

0

1

8

0

2

1

0

2

4

0

2

7

0

3

0

0

3

3

0

3

6

0

DEPTH (m)

PERCENT

STATIONS N=2171

CATCHES N=49387

SUMMER

0

5

10

15

20

25

30

35

40

45

0 3

0

6

0

9

0

1

2

0

1

5

0

1

8

0

2

1

0

2

4

0

2

7

0

3

0

0

3

3

0

3

6

0

DEPTH (m)

PERCENT

STATIONS N=278

CATCHES N=8309
Page 30

Figure 8. Mean bottom water temperature and depth at stations where juvenile and adult Atlantic herring were collected

(solid bars) and at all stations sampled (open bars) during Massachusetts inshore bottom trawl surveys, 1978-1996.

Stations

Catches

1 3 5 7 9 11 13 15 17 19 21 23

0

10

20

30

1 3 5 7 9 11 13 15 17 19 21 23

0

4

8

12

16

20

0

10

20

30

0

5

10

15

20

25

0

10

20

30

40

50

0

10

20

30

1 3 5 7 9 11 13 15 17 19 21 23

0

20

40

60

80

1 3 5 7 9 11 13 15 17 19 21 23

0

10

20

30

40

50

Juveniles Adults

Spring Spring

Spring Spring

Autumn Autumn

Autumn Autumn

Bottom Depth (m) Bottom Depth (m)

Bottom Depth (m) Bottom Depth (m)

Bottom Temperature (C) Bottom Temperature (C)

Bottom Temperature (C) Bottom Temperature (C)

Mass. Inshore Trawl Surveys

Atlantic Herring
Page 31

Figure 9a. Mean bottom water temperature and depth at stations where juvenile Atlantic herring were collected (solid

bars) and at all stations sampled (open bars) during Rhode Island Narragansett Bay trawl surveys.

-1 1 3 5 7 9 11 13 15 17 19 21 23 25 27

0

10

20

30

40

-1 1 3 5 7 9 11 13 15 17 19 21 23 25 27

0

10

20

30

40

-1 1 3 5 7 9 11 13 15 17 19 21 23 25 27

0

10

20

30

40

50

-1 1 3 5 7 9 11 13 15 17 19 21 23 25 27

0

10

20

30

40

Bottom Temperature (C)

Winter

Spring

Summer

Autumn

10 20 30 40 50 60 70 80 90 100 110 120

0

20

40

60

80

10 20 30 40 50 60 70 80 90 100 110 120

0

5

10

15

20

25

10 20 30 40 50 60 70 80 90 100 110 120 0

10

20

30

40

10 20 30 40 50 60 70 80 90 100 110 120

0

10

20

30

40

50

Winter

Autumn

Summer

Spring

Bottom Depth (ft)
Page 32

Figure 9b. Mean bottom water temperature and depth at stations where adult Atlantic herring were collected (solid bars)

and at all stations sampled (open bars) during Rhode Island Narragansett Bay trawl surveys.

-1 1 3 5 7 9 11 13 15 17 19 21 23 25 27

0

20

40

60

-1 1 3 5 7 9 11 13 15 17 19 21 23 25 27

0

3

6

9

12

15

-1 1 3 5 7 9 11 13 15 17 19 21 23 25 27

0

10

20

30

-1 1 3 5 7 9 11 13 15 17 19 21 23 25 27

0

10

20

30

40

Bottom Temperature (C)

Winter

Spring

Summer

Autumn

10 20 30 40 50 60 70 80 90 100 110 120

0

20

40

60

10 20 30 40 50 60 70 80 90 100 110 120

0

10

20

30

40

10 20 30 40 50 60 70 80 90 100 110 120

0

10

20

30

10 20 30 40 50 60 70 80 90 100 110 120 0

20

40

60

80

100

Winter

Autumn

Summer

Spring

Bottom Depth (ft)
Page 33

Figure 10a. Mean water temperature, depth, dissolved oxygen, and salinity at stations where juvenile Atlantic herring

were collected (solid bars) and at all stations (open bars) during Hudson-Raritan trawl surveys.

Temperature (C)

Depth (ft)

0 2 4 6 8 10 12 14 16 18 20 22 24 26

0

5

10

15

20

25

Stations

Catches

10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85

0

5

10

15

20

25

30

35

Dissolved Oxygen (mg/l)

Salinity (ppt)

0 1 2 3 4 5 6 7 8 9 10 11 12 13

0

5

10

15

20

25

30

35

15 17 19 21 23 25 27 29 31 33 35

0

5

10

15

20

25
Page 34

Figure 10b. Mean water temperature, depth, dissolved oxygen, and salinity at stations where adult Atlantic herring were

collected (solid bars) and at all stations (open bars) during Hudson-Raritan trawl surveys.

Temperature (C)

Depth (ft)

0 2 4 6 8 10 12 14 16 18 20 22 24 26

0

5

10

15

20

25

Stations

Catches

10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85

0

5

10

15

20

25

30

35

Dissolved Oxygen (mg/l)

Salinity (ppt)

0 1 2 3 4 5 6 7 8 9 10 11 12 13

0

10

20

30

40

50

60

70

15 17 19 21 23 25 27 29 31 33 35

0

5

10

15
Page 35

Figure 11. Distribution and abundance of Atlantic herring larvae collected during NEFSC MARMAP ichthyoplankton

surveys, January to December, 1977-1987 [see Reid et al. (1999) for details].

&OXSHDKDUHQJXV

0$50$3,FKWK\RSODQNWRQ6XUYH\V

FP%RQJR1HWPPPHVK

-DQXDU\'HFHPEHUWR

$WODQWLF+HUULQJ

1XPEHURI7RZV ZLWKODUYDH 

76 75 74 73 72 71 70 69 68 67 66 65

35

36

37

38

39

40

41

42

43

44

45

Number of Larvae / 10m2

1 to < 100

100 to < 500

500 to < 5000

5000 to < 50000

50000 to 250000

76 75 74 73 72 71 70 69 68 67 66 65

35

36

37

38

39

40

41

42

43

44

45

-DQXDU\WR

1XPEHURI7RZV ZLWKODUYDH 

0RQWKO\0HDQ'HQVLW\ /DUYDHP

Number of Larvae / 10m2

None

1 to < 10

10 to < 100

100 to 174

)HEUXDU\WR

1XPEHURI7RZV ZLWKODUYDH 

0RQWKO\0HDQ'HQVLW\ /DUYDHP

76 75 74 73 72 71 70 69 68 67 66 65

35

36

37

38

39

40

41

42

43

44

45

Number of Larvae / 10m2

None

1 to < 10

10 to 39

0DUFKWR

1XPEHURI7RZV ZLWKODUYDH 

0RQWKO\0HDQ'HQVLW\ /DUYDHP

76 75 74 73 72 71 70 69 68 67 66 65

35

36

37

38

39

40

41

42

43

44

45

Number of Larvae / 10m2

None

1 to < 10

10 to 20
Page 36

Figure 11. cont’d.

$SULOWR

1XPEHURI7RZV ZLWKODUYDH 

0RQWKO\0HDQ'HQVLW\ /DUYDHP

76 75 74 73 72 71 70 69 68 67 66 65

35

36

37

38

39

40

41

42

43

44

45

Number of Larvae / 10m2

None

1 to < 10

10 to 28

0D\WR

1XPEHURI7RZV ZLWKODUYDH 

0RQWKO\0HDQ'HQVLW\ /DUYDHP

76 75 74 73 72 71 70 69 68 67 66 65

35

36

37

38

39

40

41

42

43

44

45

Number of Larvae / 10m2

None

1 to 6

-XQHWR

1XPEHURI7RZV ZLWKODUYDH 

0RQWKO\0HDQ'HQVLW\ /DUYDHP

76 75 74 73 72 71 70 69 68 67 66 65

35

36

37

38

39

40

41

42

43

44

45

Number of Larvae / 10m2

None

1 to 7

-XO\WR

1XPEHURI7RZV ZLWKODUYDH 

0RQWKO\0HDQ'HQVLW\ /DUYDHP

76 75 74 73 72 71 70 69 68 67 66 65

35

36

37

38

39

40

41

42

43

44

45

Number of Larvae / 10m2

None

1 to 8
Page 37

Figure 11. cont’d.

76 75 74 73 72 71 70 69 68 67 66 65

35

36

37

38

39

40

41

42

43

44

45

$XJXVWWR

1XPEHURI7RZV ZLWKODUYDH 

0RQWKO\0HDQ'HQVLW\ /DUYDHP

Number of Larvae / 10m2

None

1 to < 10

10 to 70

6HSWHPEHUWR

1XPEHURI7RZV ZLWKODUYDH 

0RQWKO\0HDQ'HQVLW\ /DUYDHP

76 75 74 73 72 71 70 69 68 67 66 65

35

36

37

38

39

40

41

42

43

44

45

None

1 to < 100

100 to < 500

500 to < 5000

5000 to < 50000

Number of Larvae / 10m2

50000 to 250000

2FWREHUWR

1XPEHURI7RZV ZLWKODUYDH 

0RQWKO\0HDQ'HQVLW\ /DUYDHP

76 75 74 73 72 71 70 69 68 67 66 65

35

36

37

38

39

40

41

42

43

44

45

None

1 to < 50

50 to < 250

250 to < 1000

1000 to < 5000

Number of Larvae / 10m2

5000 to 20000

1RYHPEHUWR

1XPEHURI7RZV ZLWKODUYDH 

0RQWKO\0HDQ'HQVLW\ /DUYDHP

76 75 74 73 72 71 70 69 68 67 66 65

35

36

37

38

39

40

41

42

43

44

45

None

1 to < 50

50 to < 250

250 to < 1500

1500 to < 3000

Number of Larvae / 10m2

3000 to 7000
Page 38

Figure 11. cont’d.

'HFHPEHUWR

1XPEHURI7RZV ZLWKODUYDH 

0RQWKO\0HDQ'HQVLW\ /DUYDHP

76 75 74 73 72 71 70 69 68 67 66 65

35

36

37

38

39

40

41

42

43

44

45

None

1 to < 20

20 to < 50

50 to < 100

100 to < 250

Number of Larvae / 10m2

250 to 600
Page 39

Figure 12. Changes in abundance of Atlantic herring larvae on Georges Bank, Nantucket Shoals, and in Massachusetts

Bay from 1971-1990 (from Smith and Morse 1993). Intervals (Int.) denote periods of changing spawning patterns.
Page 40

Figure 13. Distribution and abundance of juvenile (≤ 24 cm) and adult (≥ 25 cm) Atlantic herring collected during

NEFSC bottom trawl surveys, 1963-1997. Densities are represented by dot size in spring and fall plots, while only

presence and absence are represented in winter and summer plots [see Reid et al. (1999) for details].
Page 41

Figure 13. cont’d.
Page 42

Figure 14. Distribution and abundance of juvenile (< 25 cm) and adult (≥ 25 cm) Atlantic herring collected in spring and

autumn during Massachusetts inshore bottom trawl surveys (1978-1996) [see Reid et al. (1999) for details].

Atlantic Herring

Mass. Inshore Trawl Survey

 Autumn 1978 - 1996

 Juveniles (<26cm)

Number/Tow

 1 to <25

 25 to <50

 50 to <100

 100 to <200

 200 to <512

(<25cm)

Atlantic Herring

Mass. Inshore Trawl Survey

 Spring 1978 - 1996

 Juveniles (<25cm)

Number/Tow

 1 to <10

 10 to <50

 50 to <200

 200 to <1000

 1000 to <3579

Atlantic Herring

Mass. Inshore Trawl Survey

 Autumn 1978 - 1996

 Adults (>=25cm)

Number/Tow

 1 to <5

 5 to <10

 10 to <25

 25 to <50

 50 to <213

Atlantic Herring

Mass. Inshore Trawl Survey

 Spring 1978 - 1996

 Adults (>=25cm)

Number/Tow

 1 to <2

 2 to <5

 5 to <10

 10 to <25

 25 to <58
Page 43

Figure 15. Distribution and abundance of juvenile (< 25 cm) and adult (≥ 25 cm) Atlantic herring collected in

Narragansett Bay during the Rhode Island bottom trawl survey, 1990-1996. The numbers shown at each station are the

average catch per tow rounded to one decimal place [see Reid et al. (1999) for details].

83.3

10.8

0.3

0.1

109.3

0.5

0.2

29.4

0.8

34.6

45.6

11.5 7.8

14.8

0.8

16.8

125.0

58.2

5.0

7.3

1.1

0.1

120.0

25.6

7.3 14.1

254.0

54.7

101.7

1.2

77.3

48.2

0.1

200.0

103.6

194.8

74.0

122.4 62.6

0.2

3.1

11.4

5.0

20.8

0.1

0.0

0.3

1.3

53.8

47.9

68.0 11.8

Spring

Summer Autumn

Winter

Juveniles (< 25 cm)
Page 44

Figure 15. cont’d.

3.4

8.0

0.0

2.0

1.9

0.0

0.0

2.2

0.0

11.9

11.9

5.5 0.4

1.5

0.5

0.1

2.4

0.1

0.1

0.7

0.1

0.0

4.6

1.6

3.7 0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0 0.0

0.0

0.6

0.0

0.0

2.4

0.0

0.0

0.0

0.0

9.2

0.0

0.3 8.2

Spring

Summer Autumn

Winter

Adults (> 25 cm)
Page 45

Figure 16. Distribution and abundance of Atlantic herring collected in Long Island Sound during spring and autumn

Connecticut bottom trawl surveys, 1992-1997 [see Reid et al. (1999) for details]. Histograms show lengths of herring

during spring and autumn from a subset of 1992-1997 and earlier collections.

Number/Tow

 1 to <25

 25 to <50

 50 to <100

 100 to <500

 500 to <1714

Number/Tow

 1 to <5

 5 to <25

 25 to <100

 100 to <1000

 1000 to <1328

State of Connecticut DEP

Fisheries Division

Long Island Sound Trawl Survey

SPRING 1992 - 1997

State of Connecticut DEP

Fisheries Division

Long Island Sound Trawl Survey

AUTUMN 1992 - 1997

Atlantic Herring

Atlantic Herring
Page 46

Figure 17. Distribution and abundance of juvenile (< 25 cm) and adult (≥ 25 cm) Atlantic herring collected in the

Hudson-Raritan estuary during Hudson-Raritan trawl surveys, 1992-1997 [see Reid et al. (1999) for details].

Staten

Island

NEW

YORK

NEW

JERSEY

No/Tow

 1 - 9

10 - 24

25 - 49

50 - 99

100 - 365

Atlantic Herring

Hudson-Raritan Estuary

Fall 1992-1996

Juveniles (< 25 cm)

Staten

Island

NEW

YORK

NEW

JERSEY

No/Tow

 1 - 9

10 - 24

25 - 49

50 - 99

100 - 365

Atlantic Herring

Hudson-Raritan Estuary

Winter 1992-1997

Juveniles (< 25 cm)

Staten

Island

NEW

YORK

NEW

JERSEY

No/Tow

 1 - 9

10 - 24

25 - 49

50 - 99

100 - 365

Atlantic Herring

Hudson-Raritan Estuary

Spring 1992-1997

Juveniles (< 25 cm)

Staten

Island

NEW

YORK

NEW

JERSEY

No/Tow

 1 - 9

10 - 24

25 - 49

50 - 99

100 - 365

Atlantic Herring

Hudson-Raritan Estuary

Summer 1992-1996

Juveniles (< 25 cm)
Page 47

Figure 17. cont’d.

Staten

Island

NEW

YORK

NEW

JERSEY

No/Tow

 1 - 9

10 - 24

25 - 49

50 - 99

100 - 365

Atlantic Herring

Hudson-Raritan Estuary

Fall 1992-1996

Adults ( 25 cm) >

Staten

Island

NEW

YORK

NEW

JERSEY

No/Tow

 1 - 9

10 - 24

25 - 49

50 - 99

100 - 365

Atlantic Herring

>

Hudson-Raritan Estuary

Winter 1992-1997

Adults ( 25 cm)

Staten

Island

NEW

YORK

NEW

JERSEY

No/Tow

 1 - 9

10 - 24

25 - 49

50 - 99

100 - 365

Atlantic Herring

>

Hudson-Raritan Estuary

Spring 1992-1997

Adults ( 25 cm)

Staten

Island

NEW

YORK

NEW

JERSEY

No/Tow

 1 - 9

10 - 24

25 - 49

50 - 99

100 - 365

Atlantic Herring

No Catches

Hudson-Raritan Estuary

Summer 1992-1996

Adults ( 25 cm) >
Page 48

Figure 18. Commercial landings and estimated stock biomass index (ages 2+) of Atlantic herring, 1967-1992 (Friedland

1995, 1998).

Coastal Stock Complex

Year

1965 1970 1975 1980 1985 1990 1995 2000

Metric tons (x1000)

0

500

1000

1500

2000

2500

3000

3500

4000

Commercial landings

Stock biomass (age 2+)
NORTHEAST FISHERIES SCIENCE CENTER

Dr. Michael P. Sissenwine, Science & Research Director

CAPT John T. Moakley, Operations, Management & Information Services Staff Chief

Teri L. Frady, Research Communications Unit Chief

Jon A. Gibson, Biological Sciences Editor & Laura S. Garner, Editorial Assistant

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STANDARD

 MAIL A