NOAA Technical Memorandum NMFS-NE-192

U. S. DEPARTMENT OF COMMERCE

National Oceanic and Atmospheric Administration

National Marine Fisheries Service

Northeast Fisheries Science Center

Woods Hole, Massachusetts

July 2005

Essential Fish Habitat Source Document:

Atlantic Herring, Clupea harengus,

Life History and Habitat Characteristics

Second Edition
173. Essential Fish Habitat Source Document: Barndoor Skate, Dipturus laevis, Life History and Habitat Characteristics. By

David B. Packer, Christine A. Zetlin, and Joseph J. Vitaliano. March 2003. v + 23 p., 14 figs., 1 table. NTIS Access. No. PB2003-104257.

174. Essential Fish Habitat Source Document: Clearnose Skate, Raja eglanteria, Life History and Habitat Characteristics.

By David B. Packer, Christine A. Zetlin, and Joseph J. Vitaliano. March 2003. v + 50 p., 25 figs., 2 tables. NTIS Access. No. PB2003-

104260.

175. Essential Fish Habitat Source Document: Little Skate, Leucoraja erinacea, Life History and Habitat Characteristics. By

David B. Packer, Christine A. Zetlin, and Joseph J. Vitaliano. March 2003. v + 66 p., 27 figs., 2 tables. NTIS Access. No. PB2003-104259.

176. Essential Fish Habitat Source Document: Rosette Skate, Leucoraja garmani virginica, Life History and Habitat

Characteristics. By David B. Packer, Christine A. Zetlin, and Joseph J. Vitaliano. March 2003. v + 17 p., 11 figs., 1 table. NTIS Access.

No. PB2003-104258.

177. Essential Fish Habitat Source Document: Smooth Skate, Malacoraja senta, Life History and Habitat Characteristics. By

David B. Packer, Christine A. Zetlin, and Joseph J. Vitaliano. March 2003. v + 26 p., 15 figs., 1 table. NTIS Access. No. PB2003-104256.

178. Essential Fish Habitat Source Document: Thorny Skate, Amblyraja radiata, Life History and Habitat Characteristics. By

David B. Packer, Christine A. Zetlin, and Joseph J. Vitaliano. March 2003. v + 39 p., 18 figs., 2 tables. NTIS Access. No. PB2003-104255.

179. Essential Fish Habitat Source Document: Winter Skate, Leucoraja ocellata, Life History and Habitat Characteristics.

By David B. Packer, Christine A. Zetlin, and Joseph J. Vitaliano. March 2003. v + 57 p., 25 figs., 2 tables. NTIS Access. No. PB2003-

104254.

180. Variability in Blood Chemistry of Yellowtail Flounder, Limanda ferruginea, with Regard to Sex, Season, and Geographic

Location. By Renee Mercaldo-Allen, Margaret A. Dawson, Catherine A. Kuropat, and Diane Kapareiko. September 2003. vi + 20 p., 1

fig., 10 tables. NTIS Access. No. PB2004-107074.

181. In press.

182. U.S. Atlantic and Gulf of Mexico Marine Mammal Stock Assessments – 2003. By Gordon T. Waring, Richard M. Pace, Janeen

M. Quintal, Carol P. Fairfield, and Katherine Maze-Foley, eds., Nicole Cabana, Phillip J. Clapham, Timothy V.N. Cole, Gregory L. Fulling,

Lance P. Garrison, Aleta A. Hohn, Blair G. Maise, Wayne E. McFee, Keith D. Mullin, Debra L. Palka, Patricia E. Rosel, Marjorie C. Rossman,

Frederick W. Wenzel, and Amy L. Whitingham, contribs. May 2004. vii + 287 p., 47 figs., 58 tables, 4 app., index. NTIS Access. No.

PB2004-106431.

183. In press.

184. A Large Marine Ecosystem Approach to Fisheries Management and Sustainability. By Frank J. Gable. August 2004. v +

84 p., 38 figs., 10 tables. NTIS Access. No. PB2005-101435.

185. Revised and Updated Edition of F. Bruce Sanford's 1957 "Planning Your Scientific Research Paper." By Jon A. Gibson.

August 2004. x + 36 p., 5 figs., 12 tables.

186. Essential Fish Habitat Source Document: Silver Hake, Merluccius bilinearis, Life History and Habitat Characteristics.

2nd ed. By Meredith C. Lock and David B. Packer. August 2004. v + 68 p., 28 figs., 6 tables. NTIS Access. No. PB2005-101436.

187. In preparation by authors.

188. In preparation by authors.

189. Essential Fish Habitat Source Document: Sea Scallop, Placopecten magellanicus, Life History and Habitat Characteristics.

2nd ed. By Deborah R. Hart and Antonie S. Chute. September 2004. v + 21 p., 6 figs., 2 tables. NTIS Access. No. PB2005-104079.

190. In preparation by authors.

191. Essential Fish Habitat Source Document: Northern Shortfin Squid, Illex illecebrosus, Life History and Habitat

Characteristics. 2nd ed. By Lisa C. Hendrickson and Elizabeth M. Holmes. November 2004. v + 36 p., 13 figs., 1 table. NTIS Access.

No. PB2005-101437.

Recent Issues in This Series:
U. S. DEPARTMENT OF COMMERCE

Carlos M. Gutierrez, Secretary

National Oceanic and Atmospheric Administration

Vice Admiral Conrad C. Lautenbacher, Jr., USN (ret.), Administrator

National Marine Fisheries Service

William T. Hogarth, Assistant Administrator for Fisheries

Northeast Fisheries Science Center

Woods Hole, Massachusetts

July 2005

This series represents a secondary level of scientific 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-192

David K. Stevenson1,2 and Marcy L. Scott1,3

Postal Address: 1National Marine Fisheries Serv., Northeast Regional Off., 1 Blackburn Dr.,

Gloucester, MA 01930

Essential Fish Habitat Source Document:

Atlantic Herring, Clupea harengus,

Life History and Habitat Characteristics

Second Edition

E-Mail Addresses: 2David.Stevenson@noaa.gov, 3Marcy.Scott@noaa.gov
Editorial Notes on "Essential Fish Habitat Source Documents"

Issued in the NOAA Technical Memorandum NMFS-NE Series

Editorial Production

For "Essential Fish Habitat Source Documents" issued in the NOAA Technical Memorandum NMFS-NE series, staff

of the Northeast Fisheries Science Center's (NEFSC's) Ecosystems Processes Division largely assume 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 is performed by,

and all credit for such production rightfully belongs to, the staff of the Ecosystems Processes Division.

Internet Availability and Information Updating

Each original issue of an "Essential Fish Habitat Source Document" is published both as a paper copy and as a Web

posting. The Web posting, which is in "PDF" format, is available at: http://www.nefsc.noaa.gov/nefsc/habitat/efh.

Each issue is updated at least every five years. The updated edition will be published as a Web posting only; the

replaced edition(s) will be maintained in an online archive for reference purposes.

Species Names

The NMFS Northeast Region's policy on the use of species names in all technical communications is generally to

follow the American Fisheries Society's lists of scientific and common names for fishes (i.e., Nelson et al. 2004a

; Robins

et al. 1991b), mollusks (i.e., Turgeon et al. 1998c), and decapod crustaceans (i.e., Williams et al. 1989d), and to follow the

Society for Marine Mammalogy's guidance on scientific and common names for marine mammals (i.e., Rice 1998e).

Exceptions to this policy occur when there are subsequent compelling revisions in the classifications of species, resulting

in changes in the names of species.

a

Nelson, J.S.; Crossman, E.J.; Espinosa-Pérez, H.; Findley, L.T.; Gilbert, C.R.; Lea, R.N.; Williams, J.D. 2004. Common and scientific names

of fishes from the United States, Canada, and Mexico. 6th ed. Amer. Fish. Soc. Spec. Publ. 29; 386 p. b

Robins, C.R. (chair); Bailey, R.M.; Bond, C.E.; Brooker, J.R.; Lachner, E.A.; Lea, R.N.; Scott, W.B. 1991. World fishes important to

North Americans. Amer. Fish. Soc. Spec. Publ. 21; 243 p. c

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. d

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.

e

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

PREFACE TO SECOND EDITION 

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 NOAA Fisheries to assist the 

regional fishery management councils in the 

implementation of EFH in their respective fishery 

management plans. 

NOAA Fisheries 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 a series EFH species reports (plus one 

consolidated methods report). The EFH species reports 

are a survey of the important literature as well as 

original analyses of fishery-independent data sets from 

NOAA Fisheries and several coastal states. The species 

reports are also the source for the current EFH 

designations by the New England and Mid-Atlantic 

Fishery Management Councils, and understandably are 

referred to as the “EFH source documents.” 

NOAA Fisheries 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. 

The initial series of EFH species source documents 

were published in 1999 in the NOAA Technical 

Memorandum NMFS-NE series. Updating and review 

of the EFH components of the councils’ Fishery 

Management Plans is required at least every 5 years by 

the NOAA Fisheries Guidelines for meeting the 

Sustainable Fisheries Act/EFH Final Rule. The second 

editions of these species source documents were written 

to provide the updated information needed to meet 

these requirements. The second editions provide new 

information on life history, geographic distribution, and 

habitat requirements via recent literature, research, and 

fishery surveys, and incorporate updated and revised 

maps and graphs. This second edition of the Atlantic 

herring EFH source document is based on the original 

by 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, with a foreword by Jeffrey N. Cross (Reid et 

al. 1999a). 

Identifying and describing EFH are the first steps 

in the process of protecting, conserving, and enhancing 

essential habitats of the managed species. Ultimately, 

NOAA Fisheries, 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. 
Page iv 
Page v

Contents

INTRODUCTION................................................................................................................................................................1

LIFE HISTORY ...................................................................................................................................................................1

STATUS OF THE STOCKS................................................................................................................................................6

GEOGRAPHICAL DISTRIBUTION ..................................................................................................................................7

HABITAT CHARACTERISTICS .......................................................................................................................................8

RESEARCH NEEDS .........................................................................................................................................................12

ACKNOWLEDGEMENTS................................................................................................................................................12

REFERENCES CITED – 1ST EDITION ............................................................................................................................12

NEW REFERENCES CITED – 2ND EDITION..................................................................................................................15

Tables

Table 1. Relative abundance of Atlantic herring life stages in New England and Mid-Atlantic estuaries and 

embayments by salinity zone ...........................................................................................................................19

Table 2. Distribution and abundance of juvenile Atlantic herring in New England and Mid-Atlantic estuaries and 

embayments .....................................................................................................................................................21 

Table 3. Distribution and abundance of adult Atlantic herring in New England and Mid-Atlantic estuaries and 

embayments .....................................................................................................................................................22 

Table 4. Percentage of Atlantic herring in the diets of 15 predators in the northeast U.S. Atlantic coast ecosystem ....23 

Table 5. Annual consumption (metric tons) of Atlantic herring by various predators...................................................24 

Table 6. Atlantic herring spawning site survey data and habitat parameters .................................................................25 

Table 7. Summary of habitat characteristics and requirements for Atlantic herring in the northwest Atlantic 

continental shelf ecosystem..............................................................................................................................27

Figures

Figure 1. The Atlantic herring, Clupea harengus L. (from Goode 1884)........................................................................29

Figure 2. Northeast U.S. Atlantic coast ecosystem. ........................................................................................................30 

Figure 3. Generalized view of principal Atlantic herring spawning areas in the Gulf of Maine and on Georges Bank..31 

Figure 4. Percent by weight of the major prey items in the diet of Atlantic herring. ......................................................32

Figure 5. Hypothesized seasonal movements of three Atlantic herring spawning stocks inhabiting U.S. waters...........33

Figure 6. Geo-referenced in situ observations of Atlantic herring eggs ..........................................................................34 

Figure 7. Atlantic herring spawning sites in eastern Maine, 1997-2002. ........................................................................35

Figure 8. Principal spawning grounds of Atlantic herring on Georges Bank, 1964-1971 (excluding 1967), with a 

comparison of egg patch sizes among years ....................................................................................................36 

Figure 9. Distributions and abundances of Atlantic herring larvae collected during NEFSC MARMAP 

ichthyoplankton surveys. .................................................................................................................................37 

Figure 10. Changes in abundance of Atlantic herring larvae on Georges Bank, Nantucket Shoals, and in Massachusetts 

Bay from 1971-1990 ........................................................................................................................................41 

Figure 11. Distribution of Atlantic herring larvae by age in the Georges Bank area, 1971-1990 .....................................42 

Figure 12. Distribution of recently-hatched Atlantic herring larvae on Georges Bank, 1988-1994..................................44 

Figure 13. Seasonal distributions and abundances of juvenile Atlantic herring collected during NEFSC bottom trawl 

surveys. ............................................................................................................................................................45 

Figure 14. Seasonal distributions and abundances of adult Atlantic herring collected during NEFSC bottom trawl 

surveys. ............................................................................................................................................................49 

Figure 15. Distribution and abundance of Atlantic herring along the coasts of Maine and New Hampshire....................53 

Figure 16. Length frequency plots for Atlantic herring caught along the Maine and New Hampshire coasts ..................54 
Page vi 

Figure 17. Seasonal distributions and abundances of juvenile Atlantic herring in Massachusetts coastal waters.............55 

Figure 18. Seasonal distributions and abundances of adult Atlantic herring in Massachusetts coastal waters..................57 

Figure 19. Seasonal distribution and abundance of juvenile Atlantic herring in Narragansett Bay. .................................59 

Figure 20. Seasonal distribution and abundance of adult Atlantic herring in Narragansett Bay. ......................................60 

Figure 21. Distribution and abundances of juvenile and adult Atlantic herring in Long Island Sound.............................61 

Figure 22. Relative abundance catch/tow and percent occurrence for juvenile and adult Atlantic herring in Long Island 

Sound. ..............................................................................................................................................................62 

Figure 23. Monthly log10 length frequencies (cm) of juvenile and adult Atlantic herring collected in Long Island 

Sound. ..............................................................................................................................................................63 

Figure 24. Seasonal distribution and abundance of juvenile Atlantic herring in the Hudson-Raritan Estuary..................64

Figure 25. Seasonal distribution and abundance of adult Atlantic herring in the Hudson-Raritan Estuary.......................65 

Figure 26. Catch per unit effort for total catch of juvenile and adult Atlantic herring in Chesapeake Bay, from the 

Virginia Institute of Marine Science’s (VIMS) trawl surveys..........................................................................66 

Figure 27. Seasonal distribution and abundance of juvenile Atlantic herring in Chesapeake Bay, from the VIMS trawl 

surveys .............................................................................................................................................................67 

Figure 28. Seasonal distribution and abundance of adult Atlantic herring in Chesapeake Bay, from the VIMS trawl 

surveys .............................................................................................................................................................68 

Figure 29. Atlantic herring catch per unit effort by site from the VIMS beach seine surveys. .........................................69 

Figure 30. Distributions of Atlantic herring larvae collected during NEFSC MARMAP icthyoplankton surveys relative 

to water column temperature and bottom depth ...............................................................................................70 

Figure 31. Distributions of juvenile Atlantic herring and trawls from NEFSC bottom trawl surveys relative to bottom 

water temperature, depth, and salinity..............................................................................................................71 

Figure 32. Distributions of juvenile Atlantic herring and trawls in Massachusetts coastal waters relative to bottom 

water temperature and depth. ...........................................................................................................................73 

Figure 33. Distributions of juvenile Atlantic herring in Narragansett Bay relative to mean bottom temperature and 

bottom depth. ...................................................................................................................................................75 

Figure 34. Distributions of juvenile Atlantic herring in the Hudson-Raritan Estuary relative to mean water 

temperature, depth, dissolved oxygen, and salinity..........................................................................................76 

Figure 35. Hydrographic preferences for juvenile Atlantic herring in Chesapeake Bay, from the VIMS trawl surveys ..77 

Figure 36. Distributions of adult Atlantic herring and trawls from NEFSC bottom trawl surveys relative to bottom 

water temperature, depth, and salinity..............................................................................................................78 

Figure 37. Distributions of adult Atlantic herring and trawls in Massachusetts coastal waters relative to bottom water 

temperature and depth. .....................................................................................................................................80 

Figure 38. Distributions of adult Atlantic herring in Narragansett Bay relative to mean bottom temperature and bottom 

depth.................................................................................................................................................................82 

Figure 39. Distributions of adult Atlantic herring in the Hudson-Raritan Estuary relative to mean water temperature, 

depth, dissolved oxygen, and salinity...............................................................................................................83 

Figure 40. Hydrographic preferences for adult Atlantic herring in Chesapeake Bay, from the VIMS trawl surveys. ......84 
Page 1

INTRODUCTION 

The Atlantic herring, Clupea harengus (Figure 1), 

is a pelagic, schooling, plankton-feeding species that 

inhabits both sides of the North Atlantic Ocean. In the 

western North Atlantic this species ranges from 

Labrador to Cape Hatteras and supports major 

commercial fisheries. Adult herring undergo complex 

north-south migrations for feeding, spawning, and 

overwintering. Herring produce demersal eggs and 

spawn during the summer and fall in the Gulf of Maine 

– Georges Bank region. Larvae overwinter offshore 

and in coastal waters and metamorphose into juveniles 

in the spring. Juveniles and adults are heavily preyed 

upon by a variety of marine fish, marine mammals, and 

seabirds. 

Herring are assessed and managed in U.S. waters 

as a single stock complex with two major spawning 

components, one in the Gulf of Maine and another on 

Georges Bank and Nantucket Shoals. Herring that 

spawn in the Bay of Fundy and off southwest Nova 

Scotia are assessed and managed by Canada. The U.S. 

stock complex has fully recovered from the effects of 

over-exploitation during the 1960s and 1970s and is 

currently under-utilized, although there is concern that 

exploitation rates in the Gulf of Maine may be too high. 

This report provides information on the life history, 

stock status, geographical distribution, and habitat 

characteristics of different life stages of Atlantic herring 

in U.S. and Canadian waters of the northwest Atlantic 

from Cape Hatteras to the Gulf of Maine (Figure 2). 

LIFE HISTORY 

This section provides a brief review of the biology 

and life history of Atlantic herring in U.S. and 

Canadian waters of the northwest Atlantic. More 

detailed reviews are provided by Bigelow and 

Schroeder (1953), Sindermann (1979), Kelly and 

Moring (1986), Tupper et al. (1998), and Munroe 

(2002). 

EGGS 

Atlantic herring deposit demersal eggs in 5-90 m of 

water in areas with strong tidal currents on a variety of 

substrates ranging from boulders, rocks, and gravel, to 

sand, shell fragments, and macrophytes. The eggs are 

1.0-1.4 mm in diameter (Fahay 1983) and are adhesive, 

adhering to the bottom and forming extensive egg beds 

that are often many layers deep. They remain attached 

to the bottom throughout the incubation period, which 

varies from 10-15 days in the Gulf of Maine region 

during the fall spawning season. 

LARVAE 

The larvae are 4-10 mm long at hatching (Able and 

Fahay 1998) and retain a yolk-sac for the first few days 

after hatching. The duration of the yolk-sac stage 

varies with temperature, from 2.5 days at 14.5C to 6 

days at 8C (Mansueti and Hardy 1967; Lough et al. 

1982). The yolk sac is absorbed by the time the larvae 

reach 8-12 mm SL (Blaxter and Holliday 1963). While 

they have a yolk sac, they are negatively buoyant and 

remain in deep water on or near the bottom. Divers 

observed yolk-sac larvae among the branches of a 

benthic red alga (Ptiloda serrata) at a spawning site on 

Jeffreys Ledge for several days after hatching (Cooper 

et al. 1975) and observations made from a submersible 

on Georges Bank revealed a dense aggregation of yolksac larvae being carried by the current 15 cm above the 

bottom (Caddy and Iles 1973). 

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 others are dispersed soon after hatching and 

drift with residual currents (Iles and Sinclair 1982; 

Sinclair and Iles 1985; Townsend et al. 1986; 

Chenoweth et al. 1989; Smith and Morse 1993). 

Larvae produced off southwestern Nova Scotia are 

retained initially near the spawning ground and then 

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

Power 1988). Larvae produced in coastal Gulf of 

Maine waters generally remain inshore (Graham 1982) 

and disperse in a westerly direction, entering bays and 

estuaries where they over-winter (Graham et al. 1972a; 

Graham and Townsend 1985). Some larvae are 

transported offshore, away from the coast (Townsend 

1992). Larvae that hatch on Jeffreys Ledge are 

dispersed shoreward (Boyar et al. 1973). Some larvae 

from the southwestern Gulf of Maine are transported 

eastward into estuaries in the mid-coast region of Maine 

(Lazzari and Stevenson 1992), despite the fact that the 

surface currents flow in the opposite direction. During 

the winter, herring larvae in inshore waters 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 

over-wintering in nearshore and estuarine waters or in 

coastal waters. Larvae from Nantucket Shoals and 

Georges Bank tend to drift to the southwest (Lough et 

al. 1980; Grimm 1983) and are transported as far south 
Page 2 

as southern New Jersey, where they have been collected 

in the Great Bay – Little Egg Harbor estuary during the 

winter and spring (Able and Fahay 1998) and in 

Delaware Bay and its tributaries (NEFMC 1999). 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 New England (Jury et al. 1994) and the 

Middle Atlantic (Stone et al. 1994). Larvae were 

‘highly abundant’ from Englishman-Machias Bays 

(eastern Maine) to the Sheepscot River (central Maine). 

Larvae were rare or absent in estuaries south of Raritan 

Bay (Table 1). 

The Atlantic herring is 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 and move into estuaries (Graham 1972; Fortier 

and Leggett 1983; Stephenson and Power 1988; Lazzari 

et al. 1993). 

Larvae in coastal Maine waters grow at a rate of 

about 2 mm a week between October and early January 

and from late February to March, but grow very little, if 

at all, in mid-winter (Townsend and Graham 1981). 

Lough et al. (1982) reported larval growth rates in the 

Gulf of Maine – Georges Bank area of 1.75 and 2.1 mm 

per week in the fall and less than 1 mm per week in the 

winter. 

JUVENILES 

Larvae metamorphose into juveniles at 40-55 mm 

standard length (SL) in the spring (April-May) 

(Sindermann 1979; Lough et al. 1982). Growth is 

rapid, with juveniles in coastal Maine waters reaching 

lengths of 90-125 mm by the end of their first year of 

life (Anthony 1972). Schooling behavior begins during 

metamorphosis and is well established by the time the 

larvae have made the transition to the juvenile life stage 

(Gallego and Heath 1994). In the Gulf of Maine, oneyear-old juveniles move out of nearshore waters in the 

summer and fall 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. Juvenile 

herring do not make seasonal north-south migrations. 

Herring tagged as overwintering one-year-old juveniles 

in eastern and western Maine remained in close 

proximity to the area where they were tagged 

throughout the following summer (Creaser and Libby 

1986). Some summer-feeding two-year-old juveniles 

tagged in southwestern Maine overwintered in 

Massachusetts Bay, but juveniles in eastern Maine had 

a greater tendency to remain there (Creaser and Libby 

1988). 

One and two-year-old juveniles form large schools 

in coastal waters throughout the Gulf of Maine in the 

spring and summer (Bigelow and Schroeder 1953). 

They are abundant or common in estuaries and 

embayments from Buzzards Bay to Delaware Bay, 

primarily in the spring, and have been reported in 

Narragansett Bay and Long Island Sound during all 

months of the year (Table 2). They are abundant or 

highly abundant in most estuaries and embayments 

north of Cape Cod and are particularly numerous 

between Penobscot Bay and Passamaquoddy Bay in the 

summer. 

Juvenile herring perform diurnal vertical 

migrations that are linked to changing light intensity, 

most likely in 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; activity is highest just after sunrise and 

just before sunset (Brawn 1960a; Tibbo 1964; Stickney 

1972). Juvenile and adult herring feed on a variety of 

planktonic organisms (see “Feeding”). 

ADULTS 

Like juveniles, adult herring are pelagic and form 

large schools, feeding on planktonic organisms. Adults 

in the Gulf of Maine region occupy inshore and 

offshore waters to depths of 200 m and make extensive 

seasonal migrations between summer and fall spawning 

grounds on Georges Bank and in the Gulf of Maine and 

overwintering areas in southern New England and the 

Mid-Atlantic region (see “Migrations”). Thermal 

oceanic fronts between colder and less saline 

continental shelf water and warmer, more saline 

continental slope water provide an abundance of 

plankton and other food sources and greatly influence 

the migratory behavior and spatial distribution patterns 

of this species (see “Habitat Characteristics”). 

Adults occur in estuaries and embayments from 

Passamaquoddy Bay (Bay of Fundy) to Long Island 

Sound (Table 3). They are abundant or highly abundant 

from April to November in estuaries and embayments 

north of Muscongus Bay, in mid-coast Maine. Adults 

generally are common or rare south of Long Island 

Sound. Adult herring behavior is affected by 

temperature changes. Herring probably have 

characteristic temperature ranges and tolerances during 

particular times of year (Blaxter and Holliday 1963) 

and can perceive temperature changes which are 

smaller than 0.1C (Laevastu 1993). Vertical 

migrations linked to changing light intensity are 

pronounced. Observations during the summer in the 

North Sea have shown that adult herring remain below 
Page 3

the thermocline in the daytime, migrate upwards during 

sunset to form loose aggregations above the 

thermocline, disperse between the surface and the 

thermocline during the night, and aggregate close to the 

bottom during sunrise (Laevastu and Hayes 1981; 

Blaxter and Holliday 1963). 

Median sizes and ages at maturity for male and 

female Atlantic herring during 1987-1989 autumn trawl 

surveys were 25.3 cm TL and age 2.9 years for males 

and 25.4 cm TL and 3.0 years for females (O’Brien et 

al. 1993). Boyar (1968) concluded that herring from 

the Gulf of Maine, Georges Bank, and southern Nova 

Scotia during the 1960s matured at age 4 and an 

average total length of 27.5 cm. Fewer herring matured 

at age 3 and at sizes around 26 cm. Growth and 

maturation rates appear to be density-dependant. 

Sinclair et al. (1982) correlated increased sizes at 

maturity in Nova Scotia with faster juvenile growth 

rates and Winters (1976) correlated decreased age at 

maturity with decreased adult biomass in the Gulf of St. 

Lawrence. Mean size at age of adult herring in U.S. 

waters of the northwest Atlantic has decreased steadily 

since 1983 as stock size has increased (Overholtz et al. 

2004). Growth rates have been shown to increase 

progressively from Nova Scotia to Georges Bank, with 

intermediate growth rates in the western Gulf of Maine 

(Sindermann 1979). 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 during the 1960s (Boyar 1968). 

Atlantic herring can reach a maximum size of about 39 

cm TL and 0.68 kg, and a maximum age of 15-18 years 

(Anthony 1972). However, herring caught in the U.S. 

commercial fishery seldom exceed 35 cm in length and 

12 years of age (Overholtz et al. 2004). 

REPRODUCTION 

Most Atlantic herring in the Gulf of Maine region 

mature at 3 years of age and a total length of about 25 

cm (O’Brien et al. 1993). In this report herring  25 cm 

were considered to be adults. Predicted fecundities 

range from 44,000 eggs for small (25 cm) females to 

about 250,000 for large (36 cm) females (Morse and 

Morris 1981; Kelly and Stevenson 1985). 

In the northwest Atlantic, herring spawn from 

Labrador to Nantucket Shoals. Spawning occurs in the 

spring, summer, and fall in more northern latitudes, but 

summer and fall spawning predominates in the Gulf of 

Maine-Georges Bank region (Haegele and Schweigert 

1985). Small spring spawning stocks used to exist in 

the Bay of Fundy (Bigelow and Schroeder 1953). 

Herring spawning grounds are located in high-energy 

environments with strong tidal currents (Iles and 

Sinclair 1982) and, based on information from egg and 

larval surveys and the distribution of sexually mature 

adults, are depicted - in very general terms - in Figure 

3. 

Historically, three primary herring spawning stocks 

have been recognized in the Gulf of Maine region: 

southwestern Nova Scotia, coastal Gulf of Maine, and 

Georges Bank/Nantucket Shoals. These larger stocks 

may be composed of a number of smaller stocks that 

occupy discrete, localized spawning locations within 

the larger spawning grounds. This has been confirmed 

off southwestern Nova Scotia, where spawning occurs 

on or near a series of offshore banks and ledges 

(Stephenson et al. 2001). 

In U.S. waters of the Gulf of Maine, herring eggs 

have been observed along the eastern Maine coast, at 

several other locations along the Maine coast (e.g., 

outer Penobscot Bay and near Boothbay), on Jeffreys 

Ledge and Stellwagen Bank, and on eastern Georges 

Bank (see Geographic Distribution: Eggs, and Figure 

6). Nantucket Shoals is known to be an important 

spawning ground based on the concentrations of 

recently-hatched larvae that were repeatedly collected 

there during the 1970s and 1980s (Grimm 1983; Smith 

and Morse 1993). High concentrations of recentlyhatched larvae have also been collected in the vicinity 

of Cultivator Shoals on western Georges Bank, in the 

vicinity of Stellwagen Bank and Jeffreys Ledge, and on 

the outer continental shelf in southern New England 

(Grimm 1983; Smith and Morse 1993). High densities 

of recently-hatched larvae have also been observed in 

Saco Bay and Casco Bay on the southern Maine coast 

(Graham et al. 1972b, et al. 1973). 

The spawning season in the Gulf of Maine-Georges 

Bank region begins in July and lasts until December. 

Spawning begins earlier in the northern areas of the 

Gulf. Off southwestern Nova Scotia, spawning occurs 

from July to November and peaks in SeptemberOctober (Boyar 1968; Das 1968, 1972) Spawning in 

eastern Maine coastal waters during 1983-1988 

extended from late July through early October, with 

peak spawning in late August (Stevenson 1989), but 

more recent egg bed surveys (1997-2002) in the same 

area indicated that spawning did not start until late 

August and lasted until October 21 (Neal and Brehme 

2001; Neal 2003). Based on larval surveys, Graham et 

al. (1972b) concluded that spawning peaks in midSeptember to mid-October in eastern Maine and in 

October in western Maine. Boyar et al. (1973) reported 

that spawning on Jeffreys Ledge in 1972 started in early 

September and peaked during the first three weeks of 

October. On Georges Bank, spawning occurs from late 

August to December (Boyar 1968; Berenbeim and 

Sigajev 1978; Lough et al. 1980) with a peak in 

September-October (Boyar 1968; Pankratov and 

Sigajev 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). Larval surveys conducted during 1971-

1975 indicated that spawning on Georges Bank started 

on the northeast peak of the bank in September and 
Page 4 

extended southwest to Nantucket Shoals in October, 

declined in November and was absent in December 

(Grimm 1983). 

During spawning, it has been reported that the 

females first deposit ribbons of eggs on the substrate 

and then the males swim above them and release milt 

into the water (Blaxter and Holliday 1963). However, 

Messieh (1988) observed that females spawning in 

shallow water in the Gulf of St. Lawrence did not 

release their eggs until there was milt in the water and 

swam 30 cm above the bottom for four hours before 

spawning at night. A single school of spring-spawning 

herring in a bay in southwestern Norway was observed 

to separate into a pelagic component (a tight “ball”) of 

fully-mature non-spawning fish and a demersal 

component of spawners that spread out in a flat layer at 

the bottom (Axelsen et al. 2000). Post-spawners 

seemed to return to the pelagic school. After spawning 

the two components rejoined each other in a loose, 

uneven layer at the surface. The majority of the herring 

in the school completed spawning within three days. A 

school of herring in eastern Maine was observed to 

remain near the site where they spawned for about a 

week; eggs were deposited on the bottom overnight and 

the next day the spent fish had dispersed from the 

spawning site (Stevenson and Knowles 1988). 

In some cases egg masses are composed entirely of 

eggs that were all spawned at the same time (Caddy and 

Iles 1973; Stevenson and Knowles 1988), while in other 

cases several layers of eggs in different stages of 

development indicate that successive spawnings have 

occurred at the same site within a few days of each 

other (Pankratov and Sigajev 1973). Spawning often 

occurs repeatedly at the same site. Stevenson (1989) 

reported 49 spawning events at 24 different locations 

along the eastern Maine coast during 1983-1988. In a 

few cases, eggs were deposited at the same site twice in 

the same year. 

Egg developmental rates are inversely related to 

temperature, varying from 40 days at 4-5C to 6-8 days 

at 14.4-16C, with an average incubation period of 10-

15 days at temperatures (8-13C) which prevail during 

the summer-fall spawning season in the Gulf of Maine 

(Bigelow and Schroeder 1953; Messieh 1988). In the 

Gulf of Maine, Atlantic herring spawn in fully saline 

seawater (32-33 ppt) (Munroe 2002). 

FOOD HABITS 

Atlantic herring prey upon a variety of 

planktivorous organisms. They are visual particulate 

feeders with diverse feeding behaviors, often switching 

between filtering and biting in response to light 

intensity and the size of available food (Bigelow and 

Schroeder 1953; Battle 1934; Blaxter 1966; Batty et al. 

1990). All life stages of herring are opportunistic 

feeders, and will take advantage of whatever prey of the 

appropriate size is available. As they grow and the size 

of their jaws increases, they consume larger organisms. 

Their diet therefore varies with season, their age and 

size, and location. 

Larvae begin exogenous feeding before the yolk 

sac is completely absorbed (Munroe 2002). Newlyhatched larvae (7-20 mm) in coastal waters of central 

Maine feed primarily on the small, early developmental 

stages of copepods; during the winter, larger larvae (21-

30 mm) feed on the adult stages of small copepods as 

well (Sherman and Honey 1971). During the spring, 

when a wider variety of planktonic organisms are 

available and the larvae are larger, their diet includes 

organisms such as barnacle larvae, crustacean eggs, 

copepods, and free-swimming ciliate protozoans 

(tintinnids) (Sherman and Honey 1971). Three copepod 

species preyed upon by larval herring on Georges Bank 

are 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, barnacle larvae, cladocerans, and molluscan 

larvae (Sherman and Perkins 1971). Adults have a diet 

dominated by euphausiids, chaetognaths, and copepods 

(Bigelow and Schroeder 1953; Maurer and Bowman 

1975). Maurer (1976) reported that the most important 

prey items of adult herring collected on Georges Bank 

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

euphausiids (Meganyctiphanes norvegica, 23%; 

Thysanoessa inermis, 6.1%), pteropods (Limacina 

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

Calanus finmarchicus is a common prey item. In 

addition, adults also consume fish eggs and larvae, 

including larval herring, sand lance, and silversides 

(Munroe 2002). 

Spring and summer are the most intense feeding 

times for both juvenile and adult herring (Munroe 

2002). Although it has been observed that adult herring 

on Georges Bank stop feeding prior to spawning 

(Pankratov and Sigajev 1973), there are also studies 

showing that they continue feeding before spawning 

(Bradford and Iles 1992; Axelsen et al. 2000). Feeding 

occurs primarily at dawn and dusk in the upper water 

layers due to the diurnal vertical migrations of herring 

in response to changes in light intensity; they rise to the 

surface to feed at dusk and then sink toward the seabed 

at dawn (Brawn 1960a; Tibbo 1964; Stickney 1972; 

Blaxter 1985). 

Food habits data collected during Northeast 

Fisheries Science Center (NEFSC) bottom trawl 

surveys [see Reid et al. (1999b) and Link and Almeida 

(2000)] reveal that the most abundant identifiable prey 

items (percent by weight) for Atlantic herring include 

amphipods, copepods, and euphasiids (Figure 4). 
Page 5

PREDATION AND MORTALITY 

Herring is an important species in the food web of 

the northwest Atlantic. Demersal fish species that have 

been observed feeding on herring eggs include cod, 

haddock, cunner, and red hake; invertebrates that 

probably consume herring eggs include moon snails, 

hermit crabs, and starfish (McKenzie 1964; Caddy and 

Iles 1973; Cooper et al. 1975). Herring eggs and larvae 

are consumed by sand lance (Rankine and Morrison 

1989). Herring larvae are also eaten by jellyfish 

(Aurelia aurita), Atlantic mackerel, and adult Atlantic 

herring (Bailey 1984; Bailey and Batty 1984; Moller 

1984; Lett and Kohler (1976). Juvenile herring, 

especially “brit” (age-1 juveniles) are heavily preyed 

upon due to their abundance, small size, and schooling 

behavior (Munroe 2002). 

Atlantic herring is an important prey species for a 

large number of piscivorous fish, elasmobranchs 

(sharks and skates), marine mammals, and seabirds in 

the northwest Atlantic. Unlike other pelagic fishes such 

as Atlantic mackerel, herring are smaller and vulnerable 

to predation over most, if not all, of their life (Overholtz 

et al. 2000). According to the diet composition data in 

Table 4, the principal finfish and elasmobranch species 

that feed on Atlantic herring (or on clupeid species as a 

group) are Atlantic cod, silver hake, thorny skate, 

bluefish, goosefish, weakfish, summer flounder, white 

hake, and – in certain locations and times of year – 

Atlantic bluefin tuna. Other species that feed on 

herring are spiny dogfish, Atlantic halibut, red hake, 

striped bass, dusky shark, and black sea bass. Shortfinned squid (Illex illecebrosus) have also been 

observed feeding on juvenile herring (Bigelow and 

Schroeder 1953). The spiny dogfish is a much more 

important predator on Atlantic herring than is indicated 

by diet composition data. Link et al. (2002a) estimated 

that spiny dogfish consumed an average of 67,660 

metric tons (mt) of Atlantic herring a year during 1977-

1998, with a range of 15,526 to 148,197 mt. Thus, in 

some years, spiny dogfish may consume a greater 

quantity of herring biomass than is taken in the 

commercial fishery. 

For many of the predator species listed in Table 4, 

herring made up a larger percentage of the diets of the 

larger size classes. This was the case for silver hake, 

summer flounder, white hake, bluefish, and goosefish. 

Link and Garrison (2002) reported that the percentages 

of herring in the stomachs of Atlantic cod increased 

from about 13% in 51-60 cm cod to 28% in 81-90 cm 

cod and then declined again to 6% in 111-120 cm cod. 

They also showed that herring made up a larger 

percentage of the diet of Atlantic cod in the Gulf of 

Maine than on Georges Bank or in southern New 

England. Garrison and Link (2000) reported higher 

percentages of Atlantic herring in the diet of silver hake 

on Georges Bank than in the Gulf of Maine or in 

southern New England. Bowman et al. (2000) reported 

similar results for silver hake and Atlantic cod. Chase 

(2002) reported very high percentages of Atlantic 

herring in bluefin tuna diets on Jeffreys Ledge and in 

the Great South Channel, but very low percentages in 

three other locations. Less dramatic spatial variations 

were reported for striped bass by Nelson et al. (2003). 

Overholtz et al. (2000) estimated the consumption 

of Atlantic herring by 10 species of predatory fish in 

northeastern U.S. waters from 1977-1997, and found 

that the amount of herring consumed varied in response 

to changes in the abundance of herring and the 

abundance of predator populations in the late 1980s and 

throughout the 1990s. Consumption of Atlantic herring 

by these predatory fish peaked at over 200,000 metric 

tons (mt) during 1992 and 1993, declining to less than 

100,000 mt in 1997 (Table 5). By far the most 

important predator on herring was spiny dogfish, 

followed by silver hake, cod, white hake, and bluefish. 

The declines in consumption of herring in the late 

1990s were coincident with the declines in the 

abundance of these five species. 

Read and Brownstein (2003) used survey-based 

estimates of abundance for eight species of marine 

mammals between 1991 and 1997 to estimate the total 

annual consumption of Atlantic herring by these 

species. Their estimates of marine mammal 

consumption ranged from about 94,000-190,000 mt of 

herring per year. Their results show that minke whales, 

harbor porpoises, and white-sided dolphins are major 

predators on Atlantic herring because of high 

proportions of herring (34-51%) in their diets, whereas 

fin and humpback whales consume large quantities of 

herring to sustain their large body mass. Despite a 

three-fold increase in the harbor seal population in the 

Gulf of Maine between 1981 and 1997, herring only 

make up 13% of their diet. Consequently, the mean 

consumption estimate for harbor seals is below 5,000 

mt a year. 

Read and Brownstein’s (2003) mean (or “best”) 

estimate of Atlantic herring consumed annually by 

marine mammals during 1991-1997 was about 140,000 

mt, with a range of 93,000-200,000 mt. Adding these 

estimates to the most current (1997) estimate of 

100,000 mt of Atlantic herring consumed by fish and 

elasmobranch predators reported by Overholtz et al. 

(2000) produces a total mean estimate of 240,000 mt, 

with a range of 193,000-300,000 mt. During the 1990s, 

the total amount of herring consumed by all predators 

could have been as high as 400-450,000 mt. 

MIGRATION 

Adult herring make extensive seasonal migrations 

between summer spawning grounds on Georges Bank 

and in the Gulf of Maine and overwintering areas in 

southern New England and the Mid-Atlantic region. 
Page 6 

They seldom migrate seaward beyond a depth of about 

100 m and usually inhabit waters closer to the surface 

than the bottom, except in midwinter (Hildebrand 

1963). Adults from different spawning groups 

intermingle during the non-spawning phase of their 

seasonal cycle (Sinclair and Iles 1985). Juvenile 

herring make seasonal inshore-offshore movements, but 

do not make extensive north-south migrations (see Life 

History: Juveniles). 

Three general migratory patterns are recognized off 

the northeast coast of the U.S., one for each of the three 

primary spawning stocks, based on the results of 

tagging studies (e.g., Stobo 1983 and Creaser et al. 

1984) and observations from the commercial fishery 

(Sindermann 1979; Figure 5). Herring that spawn off 

southwest Nova Scotia move north along the eastern 

Scotian shelf after spawning, but some also move south 

to overwinter in the Gulf of Maine. Adults belonging 

to the Georges Bank/Nantucket Shoals stock overwinter 

south of Cape Cod and along the Mid-Atlantic coast. 

They move 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. Adults that spawn in the Gulf of Maine 

migrate southwest along the coast after spawning. 

Some of them overwinter south of Cape Cod and some 

remain in the southwestern region of the gulf. Thermal 

oceanic fronts between colder, and less saline 

continental shelf water and, warmer, more saline 

continental slope water provide an abundance of 

plankton and other food sources and greatly influence 

the migratory behavior of this species (see Habitat 

Characteristics: Adults). 

STATUS OF THE STOCKS 

Adult herring segregate into discrete spawning 

stocks in the summer and fall – on Georges Bank and 

Nantucket Shoals, in coastal waters of the Gulf of 

Maine, and off southwest Nova Scotia and in the Bay of 

Fundy. Each of the major spawning areas in the Gulf of 

Maine region consists of a number of smaller, discrete, 

spawning sites. Some degree of stock differentiation 

was achieved with early enzyme electrophoresis 

research (Ridgway et al. 1970, 1971), but more recent 

attempts to differentiate geographically isolated fall 

spawning stocks in eastern Canada and the northeast 

U.S. on the basis of more specific genetic 

characteristics have been unsuccessful (Kornfield et al. 

1982; Kornfield and Bogdanowicz 1987; Safford and 

Booke 1992). Evidence for separate stocks is based on 

discrete larval distribution patterns (Iles and Sinclair 

1982), differences in spawning times and locations 

(Boyar et al. 1973; Haegele and Schweigert 1985), 

distinct biological characteristics - such as growth rates 

(Anthony and Waring 1980) and meristic and 

morphometric characteristics (Anthony 1981; Safford 

1985) - and the incidence of parasites (McGladdery and 

Burt 1985). McQuinn (1997) reviewed arguments for a 

discrete versus dynamic balance population concept for 

Atlantic herring and 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. 

The most compelling evidence supporting the 

existence of separate Gulf of Maine and Georges BankNantucket Shoals stocks was the collapse of the large 

Georges Bank-Nantucket Shoals stock in the early 

1970s after several years of heavy exploitation by 

foreign fishing fleets (Overholtz and Friedland 2002). 

This stock remained in a depleted state for 10-15 years, 

during which time the smaller Gulf of Maine stock 

continued to support a strong coastal fishery. 

Trawl and larval survey data show that the Georges 

Bank stock has fully recovered and support the view 

that herring recolonized the bank in stages from the 

Gulf of Maine and Nantucket Shoals during the late 

1980s (Smith and Morse 1993; Overholtz and Friedland 

2002). Analysis of trawl survey shows that the 

geographic range of herring in U.S. waters of the 

northwest Atlantic was greatly reduced during the 

period of stock depletion and was more widely 

dispersed by the mid 1990s (Overholtz 2002; Overholtz 

and Friedland 2002). During 1968-1970 the springtime center of distribution was south and west of Cape 

Cod and then gradually shifted northwards and 

eastwards as stock size declined. As abundance 

increased in the late 1980s, the center of the spring 

distribution moved southwards and westwards as adults 

that spawn on Georges Bank and on Nantucket Shoals 

migrated south and re-occupied the mid-Atlantic shelf. 

The Bay of Fundy-southwest Nova Scotia stock is 

assessed by Canada as a component of a larger 

management unit that also includes coastal and outer 

shelf waters east of Nova Scotia. Biomass estimates 

derived from acoustic survey data indicate that the 

overall abundance of spawning herring declined from 

about 570,000 mt in 1997 to about 460,000 mt in 2000 

and 2001, but increased in 2002 and 2003 (Melvin et al. 

2004). Despite recent increases in spawning stock 

biomass (SSB), there are concerns that the stock is 

deteriorating: the total catch increased in 2003, but 

there are fewer adults in the population and SSB for 

two spawning components remains well below 

historical levels (Power et al. 2004). The abundance of 

herring that spawn at individual sites off southwest 

Nova Scotia varies from site to site in response to the 

amount of fishing that occurs at each site (Stephenson 

et al. 1999; Melvin et al. 2001). These observations 

support the view that each of these spawning 

aggregations constitutes a separate sub-stock of herring 

(Stephenson et al 2001). Some of these discrete 
Page 7

spawning sites are located within 10-15 miles of each 

other. 

Herring that spawn on Georges Bank, Nantucket 

Shoals, and in coastal waters of the Gulf of Maine are 

currently assessed in the U.S. as a single coastal stock 

complex. According to a recent U.S. assessment, 

spawning stock biomass for the stock complex was 

about 1.4 million metric tons (mt) in 2001 while a 

Canadian assessment shows it to be about 600,000 mt 

(Overholtz et al. 2004). They both show the same 

downward trend in spawning stock size from about one 

million metric tons in the late 1960s to 100,000 mt 

between 1975 and 1985, but the U.S. assessment 

indicates a much more dramatic recovery during the last 

20 years. Maximum sustainable yield (MSY) estimates 

from the U.S. assessment were 222,000 mt or 243,000 

mt, based on two different model formulations 

(Overholtz et al. 2004). According to the U.S. 

assessment, current fishing mortality rates in the fishery 

are below 10%, indicating that the resource is 

significantly under-utilized. There is concern, however, 

that the inshore (Gulf of Maine) component of the 

stock, which is heavily exploited, is being overharvested. 

GEOGRAPHICAL DISTRIBUTION 

EGGS 

Atlantic herring eggs are demersal and adhere to 

the substrate and were not usually collected during the 

NEFSC Marine Resources Monitoring, Assessment and 

Prediction (MARMAP) ichthyoplankton surveys. A 

few in situ surveys of herring spawning locations have 

been conducted in the Gulf of Maine – Georges Bank 

region during the past 40 years using divers, remotelyoperated underwater vehicles equipped with video 

cameras, submersibles, dredges and grab samplers. 

Information obtained from these surveys is summarized 

in Table 6. Geo-referenced spawning site locations are 

shown in Figure 6, Figure 7, and Figure 8. Spawning 

location information in this document is based on the 

presence of herring eggs on the bottom. More general 

information on herring spawning grounds based on 

catches of fully mature adults or the abundance and 

distribution of recently-hatched larvae is summarized in 

Life History: Reproduction. 

LARVAE 

Herring larvae were collected during the 1977-

1987 NEFSC MARMAP surveys from New Jersey to 

the Bay of Fundy and from nearshore waters to the 

seaward limit of the survey area (Figure 9). Larvae 

were collected in all months, but were most abundant in 

the fall (September – December). The highest mean 

monthly density (351 larvae/10 m2

) occurred in 

September 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

). The appearance of larvae off Nova 

Scotia and eastern Maine in September, followed by 

more widespread larval production throughout the Gulf 

of Maine and on Nantucket Shoals and Georges Bank 

in October and November, indicates that spawning 

begins earlier in the northeast (see also Bigelow and 

Schroeder 1953; Tupper et al. 1998). Mean densities 

were much lower (less than 6 larvae/10m2

) from 

December through August. The MARMAP surveys 

were conducted during the time when the abundance of 

the Georges Bank – Nantucket Shoals spawning stock 

of herring was very low. 

The abundance and distribution of herring larvae 

on Georges Bank and Nantucket Shoals changed 

considerably between 1971 and 1990, as the number of 

herring spawning in these two areas declined in 

response to heavy fishing pressure and then increased 

beginning in the mid-1980s. By the end of the 1980s, 

larval abundance had increased to 1973-1974 levels on 

Georges Bank and exceeded the previous abundance 

levels on Nantucket Shoals (Figure 10). According to 

U.S. larval survey data compiled by Smith and Morse 

(1993), herring spawned on the northeast peak of 

Georges Bank and on Nantucket Shoals during 1971-75 

(Figure 11). During 1976-1987, spawning was limited 

to a small area on Nantucket Shoals and in 

Massachusetts Bay. During 1988-1990, spawning 

spread over a larger area that included the western 

portion of Georges Bank (Cultivator Shoals), but not 

the northeast peak. More recent Canadian surveys have 

documented larval production in U.S. and Canadian 

waters on eastern Georges Bank, including – in 1994 – 

the northeast peak (Figure 12; Melvin et al. 1996). 

JUVENILES AND ADULTS 

NEFSC bottom trawl surveys show that the 

distributions of juvenile (Figure 13) and adult (Figure 

14) herring overlap during the summer, fall, and winter 

and are very similar in the spring. (Note that winter and 

summer distributions are presented as presence/absence 

data, precluding a discussion of abundances.) In the 

summer and fall, herring are distributed throughout the 

Gulf of Maine and in the deeper waters of Georges 

Bank and the Great South Channel, with a few in 

offshore waters of southern New England. In the 

winter, their distribution shifts southward, extending 

from Georges Bank to Cape Hatteras, primarily in 

offshore waters. A few remain in the Gulf of Maine, 
Page 8 

which is not sampled very heavily in the winter (Reid et 

al. 1999b). In the spring, juvenile herring occupy the 

entire region. The spring adult distribution extends 

from Cape Hatteras to Georges Bank and the 

southwestern and central Gulf of Maine and the Scotian 

shelf, but there are only a few along the Maine coast. 

Herring are more concentrated in nearshore waters of 

the Mid-Atlantic, southern New England, and 

Massachusetts Bay in the spring than they are in the 

winter. 

The distributions and abundances of Atlantic 

herring along the coasts of Maine and New Hampshire, 

based on spring and fall 2000-2003 Maine-New 

Hampshire inshore groundfish surveys (Sherman et al. 

2004), are shown in Figure 15. Most of these were 

juveniles (Figure 16). 

The distributions and abundances of juveniles and 

adults in Massachusetts coastal waters, based upon the 

spring and fall 1978-2003 Massachusetts inshore trawl 

surveys, are shown in Figure 17 (juveniles) and Figure 

18 (adults). In the spring, the largest catches of juvenile 

herring occurred along the northern shore of Nantucket 

Island and southern shore of Martha's Vineyard, as well 

as in Buzzards Bay north of Cape Ann. In the fall, large 

catches were found in Cape Cod Bay (Figure 17). The 

few adults found in the spring and fall were most 

abundant in near Cape Ann (Figure 18). 

The seasonal distributions and abundances of 

juveniles and adults in Narragansett Bay, based upon 

the 1990-1996 Rhode Island bottom trawl surveys, are 

shown in Figure 19 (juveniles) and Figure 20 (adults). 

Catches of juveniles were patchy in Narragansett Bay 

(Figure 19). 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 (Figure 20) 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. 

The distributions and abundances of both juvenile 

and adult Atlantic herring in Long Island Sound from 

April to November 1984-1994, based on the 

Connecticut Fisheries Division bottom trawl surveys, 

are shown in Figure 21, Figure 22, and Figure 23. The 

following description of their distributions relative to 

depth and bottom type is taken verbatim from 

Gottschall et al. (2000). 

Atlantic herring taken in the survey ranged from 3-

33 cm (Figure 23). The percentage of adults was 81% 

and 94% in April and May respectively, but declined to 

0% by October. Although herring were not measured in 

November, recorded observations indicated that most 

were adults (Gottschall et al. 2000). 

Atlantic herring abundance in the Connecticut 

survey was highest during April and May (Figure 22) 

when adults were most abundant in Long Island Sound. 

In the spring sampling period they were widely 

distributed, but were especially abundant in the Western 

and Central Basins over mud bottom (Figure 21 and 

Figure 22B). Adult abundance declined through the 

spring months to very low abundance in the summer 

and fall periods, when most of the herring taken in the 

survey were juveniles. Although the survey did not 

effectively retain young-of-the-year, they were very 

abundant in the Sound during the summer months – in a 

separate sampling program, up to 80,000 per 15 minute 

tow were caught with an otter trawl equipped with a 6 

mm codend liner [see reference in Gottschall et al. 

(2000)]. During the fall period, most herring were taken 

along the Connecticut side of the Sound in depths < 18 

m, especially south of Milford (Figure 21 and Figure 

22F). Abundance increased in November when adult 

fish were again taken. During November, abundance 

increased slightly with decreasing depth, and the largest 

catches occurred in the Central Basin (Gottschall et al. 

2000). 

Surveys of the Hudson-Raritan estuary show that 

juveniles were most abundant in winter and spring 

throughout the (Figure 24). Some were caught at the 

mouth of the estuary in summer, but they were rare in 

the fall. Adults (Figure 25) were common in winter, 

but not at any other time of year. 

The Virginia Institute of Marine Science (VIMS) 

trawl surveys (1988-1999) and beach seine surveys 

(1994-1999) of Chesapeake Bay show that, although 

Atlantic herring were not often caught, nearly 90% of 

the catches were juveniles, with only three adults found 

from the seine surveys (Geer 2002). Juveniles were 

caught in the trawl survey during late winter and early 

spring, with peaks in April and May (Figure 26). 

During the winter juveniles were caught mostly in the 

tributaries but were distributed throughout the lower 

Chesapeake Bay in the spring (Figure 27). In the 

summer a few juveniles were found at the mouth of the 

Bay, and in the fall they were found in the mainstem of 

the Bay. Adults were only caught in the trawl survey 

during the winter (Figure 26 and Figure 28). Of the 

9,321 herring captured during the beach seine surveys, 

over 86% came from a single sample in May 1996 

(Geer 2002). They were found at the Bay mouth or 

along the Atlantic coast beaches (Figure 29). 

HABITAT CHARACTERISTICS 

Information on the habitat characteristics of 

Atlantic herring are presented here and summarized in 

Table 6 and Table 7. 
Page 9

EGGS 

Atlantic herring eggs are spawned on the bottom in 

discrete beds in coastal and offshore waters of the Gulf 

of Maine, Georges Bank, and Nantucket Shoals. 

Depths reported during in situ egg bed surveys ranged 

from 5 m at Grand Manan Island to 73 m at a spawning 

site in eastern Maine (Table 6). Eggs have been 

reported on Jeffreys Ledge between depths of 53 and 59 

m. Pre-spawning aggregations of adult herring have 

been observed on the northern edge of Georges Bank in 

depths of 50-100 m (Pankratov and Sigajev 1973). 

Munroe (2002) reported the maximum spawning depth 

in the Gulf of Maine region to be about 90 m. 

Eggs have been observed on a variety of substrates 

that include rocks (ranging from pebbles to boulders), 

gravel, shell fragments, sand, and benthic macroalgae 

and epifauna attached to hard substrates (Table 6). 

Spawning sites have also been located by interviewing 

lobster fishermen who have seen eggs attached to their 

traps (Stevenson 1989). Gravel and shell fragments 

have been identified as the preferred substrate for 

herring eggs in nearshore spawning areas in eastern 

Maine, gravel and rocks with an attached red alga on 

Jeffreys Ledge, and gravel on Georges Bank. Fine sand 

and mud are not good substrates for herring eggs and 

often define the edges of egg beds. Fine sediments do 

not provide a stable substrate for attached eggs 

(Drapeau 1973) and are not characteristic of relatively 

shallow, tidally-energetic benthic habitats where 

herring spawn (Iles and Sinclair 1982). Egg beds on 

the northern edge of eastern Georges Bank surveyed by 

U.S. and Soviet scientists between 1964 and 1970 were 

all located in elongated ridges of gravel at depths of 40-

50 m between gravelly sand and large sand ridges 10-

20 m in height (Valentine and Lough 1991). 

In the Gulf of Maine region, herring eggs have 

been observed as individual eggs, clumps, or patches 

and in cohesive mats up to 5 cm thick (Table 6). In 

some cases egg beds are quite uniform in thickness and 

in others they vary considerably, from several 

centimeters deep to individual eggs at the edges of the 

egg bed (Table 6). Egg mortality is directly related to 

current speed and the amount of oxygen that is 

available to eggs in the lower layers. Stevenson and 

Knowles (1988) observed that eggs throughout an egg 

mass 3 cm thick developed at the same rate and that in 

most samples collected from the egg mat, less than 1% 

of the eggs were dead (egg mortality was less than 5% 

in all samples). In contrast, Cooper et al. (1975) 

observed 50-70% egg mortality in the lower portion of 

an egg mat 4-5 cm thick. 

The sizes of egg beds that have been surveyed in 

the Gulf of Maine and on Georges Bank have varied 

from 0.07-1.39 km2

 (Table 6). Egg beds on the eastern 

Maine coast are typically longer than they are wide, 

following depth contours that parallel the shoreline 

(Figure 7). Egg beds that were surveyed on offshore 

banks (Georges Bank and Trinity Ledge) were more 

irregular in shape (McKenzie 1964; Pankratov and 

Sigajev 1973). Eggs on Georges Bank, Trinity Ledge, 

and eastern Maine were all deposited in fairly flat or 

gradually sloping bottom areas. Eggs at one site on 

Jeffreys Ledge were found on top of an underwater 

“hill” and down a 20-40 degree rocky slope to talus 

material at the base of the hill and beyond (Cooper et 

al. 1975). 

Herring spawning sites are characterized by strong 

bottom currents. Tidal currents up to two knots in 

velocity have been measured during egg bed surveys in 

the Gulf of Maine and on Georges Bank (Table 6). 

Strong currents prevent siltation (which would impede 

egg adherence to the substrate and smother eggs), 

supply oxygen to the developing eggs, and remove 

metabolites (Drapeau 1973). 

Bottom temperatures measured during herring egg 

bed surveys in the Gulf of Maine region have ranged 

from a low of 7C to a high of 15C (Table 6). The 

temperature range for normal egg development and 

survival is not known with certainty. Bigelow and 

Schroeder (1953) reported that temperatures above 

20C and below 5C were lethal, but experiments in 

Europe have shown that egg development was normal 

between 1C and 22C, with mortality at –0.8C 

(Blaxter and Holliday 1963). Slightly lower minimum 

temperatures have ranged from –1.2 to 0C for herring 

stocks off northern Europe (Kelly and Moring 1986). 

Atlantic herring in the Gulf of Maine spawn in 

fully saline seawater (32-33 ppt) (Munroe 2002). 

However, laboratory experiments show that 

fertilization, egg development, and hatching can 

succeed in salinities of 5.9-52.5 ppt, with maximum 

fertilization at 25 ppt or more and maximum hatching 

success at 20-35 ppt (Holliday and Blaxter 1960). 

Hatching success is low at dissolved oxygen 

concentrations below 20% saturation (Bishai 1960). 

Eggs covered with 1 cm of sediment do not survive 

(100% mortality) while a thin film of sediment causes 

85% mortality, but there was no effect of suspended 

sediments at any concentration up to 7,000 mg/l on 

hatching success (Messieh et al. 1981). Eggs incubated 

in 30 micrograms/l copper during incubation had 

relatively high mortality rates and premature hatching, 

with 70% of the larvae being deformed (Blaxter 1977). 

LARVAE 

Once the yolk-sac is absorbed (within a few days 

after hatching), herring larvae are pelagic and begin to 

feed on planktonic organisms. They are transported 

away from spawning areas and overwinter in inshore 

bays and estuaries, or in offshore coastal waters, for 4-8 

months before metamorphosing to juveniles in the 

spring. Herring larvae are tolerant of wide ranges of 
Page 10

temperature and salinity, and low oxygen 

concentrations. In laboratory experiments, upper and 

lower temperature tolerances of newly hatched herring 

larvae were 22-24C and –0.75 to –1.8C (Blaxter and 

Holliday 1963). Larvae tolerated salinities of 1.4-60 ppt 

for 24 hours and 2.5-52.5 ppt for 7 days (Blaxter and 

Holliday 1963; Blaxter and Hunter 1982). De Silva and 

Tytler (1973) reported 50% mortality of larvae exposed 

to oxygen concentrations of 1.9-3.6 mg/l at 10C after 

96 hours. Eggs and larvae held under films of crude oil 

in concentrations of 1 ml or 20 ml/l or in emulsions 

experienced toxicities that varied with the origin of the 

oil (Kühnhold 1969, as cited in Blaxter and Hunter 

1982). Fractions with lower boiling points seemed more 

harmful. Larvae swim into oil dispersants and are 

narcotized (Wilson 1974). High mortality of newly 

hatched larvae has been observed at copper 

concentrations of 1,000 micrograms/l; larvae were more 

resistant than eggs (Blaxter 1977). 

In the NEFSC MARMAP survey, most larvae were 

collected at 8-14°C from September to November; 

maximum abundance was at 9-12°C (Figure 30). In 

December, larvae occurred at 6-11°C with the majority 

collected at 8-9°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. 

JUVENILES 

Laboratory experiments have shown that juvenile 

herring tolerate higher and lower temperatures than 

adults. The preferred temperature range for juveniles is 

8-12°C and physiological stress has been observed at 

temperatures below 4°C and above 16°C (Stickney 

1969). Brawn (1960b) reported that 50% of juvenile 

herring exposed to temperatures between 19.5 and 

21.2°C died within 48 hours and that they survived at 

temperatures as low as -1.1°C. The blood of Atlantic 

herring contains antifreeze proteins (AFP) which allows 

them to survive in icy seawater. Plasma-freezing points 

are significantly lower and AFP activity significantly 

higher in juveniles than in adults (Chadwick et al. 

1990). 

Salinity is probably not as important a factor as 

temperature in affecting the distribution and movements 

of Atlantic herring (Munroe 2002). There is a tendency 

for herring to prefer higher salinities and to avoid 

brackish conditions with increasing age. Laboratory 

studies indicate that juveniles prefer salinities of 28-32 

ppt (Stickney 1969), and can tolerate salinities as low as 

5 ppt for brief periods of time (Brawn 1960c). Their 

salinity preference is temperature dependent. Stickney 

(1969) reported that juveniles preferred salinities above 

29 ppt at temperatures below 10°C, but there was no 

salinity preference at temperatures above 10°C. 

Juveniles occupy inshore coves and estuaries with low 

salinities in the spring and summer of their first year of 

growth (Townsend 1992), whereas older juveniles 

avoid brackish estuarine conditions (Recksieck and 

McCleave 1973). 

The spring and fall distributions of juvenile 

Atlantic herring relative to bottom water temperature, 

depth, and salinity based on NEFSC bottom trawl 

surveys from the Gulf of Maine to Cape Hatteras are 

shown in Figure 31. In the spring, juveniles were found 

between 2-12°C, with most between 3-7°C. During 

autumn, they were found between 5-17°C, with the 

majority between 6-10°C. They occurred on the outer 

continental shelf to a maximum depth of 300 m in the 

spring and fall. In the spring, the majority were found in 

depths < 100 m. They were caught in a salinity range of 

30-35 ppt in the spring and 32-35 ppt in the autumn, 

with the majority found at 32-33 ppt during spring and 

33-34 ppt in the autumn. 

The spring and autumn distributions of juvenile 

Atlantic herring in Massachusetts coastal waters 

relative to bottom water temperature and depth based 

on 1978-2003 Massachusetts inshore trawl surveys are 

shown in Figure 32. Juveniles were collected at 

temperatures ranging from 1-16°C during the spring 

and 4-21°C during the autumn. Catch rates were high at 

11°C in the spring and at 8°C in the fall. In the spring, 

they were found over depths ranging from 6-85 m, with 

higher catches between 11-20 m. During autumn they 

were found from 1-85 m, with a very high catch at 21-

25 m. 

In the Narragansett Bay bottom trawl survey, most 

juveniles were caught at 17-21°C in the summer, and 

10-11°C and 18-20°C in the fall, 2-6°C in the winter, 

and 10 and 12°C in the spring (Figure 33). Most were 

caught between depths of 11-40 ft (3-12 m) in the 

spring, summer, and fall surveys. Catch rates were high 

at bottom depths of 91-100 ft (27-30 m) in all seasons 

and also at 51-60 ft (15-18 m) in spring. 

The distributions and abundances of both juvenile 

and adult Atlantic herring in Long Island Sound relative 

to depth and bottom type were discussed previously 

under Geographic Distribution: Juveniles and Adults 

(Figure 22; Gottschall et al. [2000]). 

In the Hudson-Raritan estuary, most juvenile 

herring were caught during winter, spring, and summer 

bottom trawl surveys at 3-5°C and 14-21°C, but they 

were most abundant at 15-18°C (Figure 34). Most were 

caught between 15 ft and 55 ft (4-16 m), dissolved 

oxygen (DO) concentrations of 6-12 mg/l, and salinities 

of 21-31 ppt. Catch rates were highest in depths of 30-

55 ft (9-16 m), DO levels of 10 and 11 mg/l, and 

salinities of 21, 26, 27 and 31 ppt. Most juveniles were 

caught in the winter and spring surveys, with some in 

the summer and very few in the fall (Figure 34). 

The hydrographic preferences of juvenile Atlantic 

herring in Chesapeake Bay from the 1988-1999 VIMS 
Page 11

trawl surveys are shown in Figure 35 (all years and 

months combined). Geer (2002) states that the juveniles 

are found primarily at temperatures between 10-16°C 

and at salinities > 14 ppt. It appears they prefer 

dissolved oxygen levels of 9 mg/l and depths of < 10 m 

(Figure 35). 

ADULTS 

Like juvenile herring, adults utilize pelagic 

habitats, only using the bottom for spawning. 

Observations of seasonal distribution on Georges Bank 

suggested a preferred temperature range of 5-9°C 

(Zinkevich 1967). Adults regularly enter bays and 

estuaries, but are rarely found in low salinities 

(Hildebrand 1963; Munroe 2002). Bigelow and 

Schroeder (1953) reported that the lower limit of 

salinity for adult Atlantic herring in the Gulf of Maine 

was probably 28 ppt. 

Factors which may affect herring distribution 

include currents and frontal zones (Sutcliffe et al. 1977; 

Sinclair and Iles 1985). Jakobsson (1980) reported that 

the densest concentrations of herring in Icelandic 

waters were in waters just at or just inside the edge of 

the continental shelf in boundary areas of warm and 

cold water masses. Depth, substrate type, and 

zooplankton abundance were significant factors 

affecting the presence and relative abundance of adult 

herring in the northern North Sea in 1992, 1994, and 

1995 (Maravelias 1999). Herring were more abundant 

in depths < 150 m, on plankton “patches” or on their 

edges, and on gravel-sand seabeds where they spawned 

1-2 months later. These relationships were stable over 

time, despite a substantial reduction in stock size 

(Maravelias et al. 2000b). Further analysis showed that 

there were more herring in areas where sea surface 

temperatures were between 11°C and 14°C, the 

thermocline was 25-45 m deep, and the difference 

between surface and bottom waters was only 3°C 

(Maravelias et al. 2000a). In more stratified waters 

(with colder bottom water), herring abundance 

decreased. This research supports the hypothesis that 

well-mixed waters and transition zones between wellmixed and stratified waters are preferred habitats for 

adult herring. Furthermore, as stock size decreased, 

herring aggregated in fewer, more distinct regions with 

these habitat characteristics. Schools were also found 

preferentially over areas of hard substrate and there was 

a strong relationship with particular topographic 

features within the survey area; i.e., a low ridge and two 

escarpments (Reid and Maravelias 2001). 

Fronts created by currents and eddies act as 

distribution boundaries for herring through their direct 

effects on the fish themselves, and also indirectly by 

aggregating planktonic food organisms and increasing 

the production of zooplankton. Mixing, such as occurs 

in the frontal zones at the edge of the continental shelf 

or as a result of increased current flow and turbulence 

on the edges of offshore banks causes elevated nutrient 

levels, increased primary production, and increased 

zooplankton abundance. 

The spring and fall distributions of adult Atlantic 

herring relative to bottom water temperature, depth, and 

salinity based on NEFSC bottom trawl surveys from the 

Gulf of Maine to Cape Hatteras are shown in Figure 36. 

In the spring, adults were found between 2-13°C, with 

most between 4-7°C. During autumn, they were found 

between 4-16°C, with the majority between 6-10°C. 

They occurred on the outer continental shelf to a 

maximum depth of 300 m in the spring and the fall. In 

the fall, most were found > 80 m, while in the spring, 

the majority were found at shallower depths. They were 

caught in a salinity range of 27-35 ppt in the spring and 

32-35 ppt in the autumn, with the majority found at 33 

ppt during spring and 33-34 in the autumn. 

The spring and autumn distributions of adult 

Atlantic herring in Massachusetts coastal waters 

relative to bottom water temperature and depth based 

on Massachusetts inshore trawl surveys are shown in 

Figure 37. Adults were collected at temperatures 

ranging from 2-13°C during the spring and 4-14°C 

during the autumn. Most were found at 4-5°C in the 

spring and at 6-10°C in the fall; the catch was high at 

7°C in the fall. In the spring, they were found over 

depths ranging from 6-85 m, with higher catches 

between 46-50 m and 76-80 m. During autumn they 

were found from 21-85 m, with most at depths > 50 m. 

In Narragansett Bay during the winter, most adults 

were caught in bottom temperatures of 3-7°C and 

depths of 20, 40-60, and 100 ft (6-30 m), while in the 

spring, most were caught at 3-5°C and 7-11°C and 

depths of 20-40, 70 and 100 ft in the spring (Figure 38). 

High catch rates occurred at 6°C in winter and 4-5°C in 

spring and at 100 ft at both times of year. In the fall, 

high catches occurred at 12°C and at 30 ft. Most adults 

were caught in the winter and spring surveys: none 

were caught in the summer. 

In the Hudson-Raritan estuary, most adults were 

caught at bottom temperatures of 2-6°C, depths of 15-

45 ft (4-14 m), dissolved oxygen (DO) concentrations 

of 9-12 mg/l, and salinities of 24-33 ppt. Catches were 

highest at 3-6°C, depths of 15-25 ft (10-14 m), DO 

levels of 11 mg/l, and salinities of 24-25 and 28-31 ppt. 

Most adults were caught in the winter (Figure 39). 

The hydrographic preferences of the few adult 

Atlantic herring caught in Chesapeake Bay during the 

1988-1999 VIMS trawl surveys are shown in Figure 40 

(all years and months combined). Adults were found in 

only during the winter at greater depths and colder 

waters than the juveniles (Geer 2002). They preferred 

dissolved oxygen levels of 11 mg/l and salinities > 14 

ppt. 
Page 12

RESEARCH NEEDS 

The following research needs are based in part on a 

summary provided by Tupper et al. 1998. 

Discrete populations/metapopulations within the 

Atlantic coastal stock complex need to be identified. 

This would involve identifying the major and minor 

spawning components within the Gulf of Maine – 

Georges Bank region and the degree to which they 

intermix at different times of year. Research methods 

that should be considered include the examination of 

environmentally-induced traits (scales, otoliths, and 

possibly morphometrics), and tagging studies. Tagging 

studies could be conducted in conjunction with other 

surveys (e.g., acoustic surveys during the summer and 

fall spawning seasons). Modern genetic techniques 

(e.g., cDNA fingerprinting) and physiological 

performance indices might be useful for stock 

differentiation purposes and should be evaluated. 

Pertinent questions that should be addressed include: 

 During the winter, what is the degree of 

mixing among adults that spawn on Georges 

Bank, Nantucket Shoals, and in the Gulf of 

Maine and migrate to southern New England 

and the mid-Atlantic region? 

 During the summer and fall, what is the degree 

of mixing between adults that spawn in 

different locations? 

Given the concerns that have been expressed 

regarding the status of the Gulf of Maine spawning 

stock (see Status of The Stocks), stock assessment 

surveys or analyses that would indicate trends in 

population size in the Gulf of Maine are badly needed. 

Procedures used to estimate population size or resource 

status for the different components of the coastal stock 

complex are complicated by the fact that adults from 

each spawning group mix to an unknown degree in 

different geographical areas and at different times of 

year. A large-scale larval herring survey, conducted 

repeatedly (e.g., at two-week intervals) throughout the 

spawning season on all known spawning grounds, 

would provide useful information for comparing the 

relative intensity of spawning by the different 

components of the resource and for evaluating to what 

extent larval production (and spawning stock size) on 

Georges Bank, Nantucket Shoals, and the southwest 

Gulf of Maine has changed since the last large-scale 

larval herring survey was done in these areas in 1990. 

Existing surveys of pre-spawning aggregations of 

adult herring should be continued: currently, acoustic 

surveys are conducted in the Gulf of Maine and on the 

northern edge of Georges Bank (See Overholtz et al.

2004). New technologies, such as multi-beam acoustic 

equipment, towed-array video, and laser illumination, 

could provide surveying tools. The applicability of 

alternative assessment models that have been used with 

other pelagic resource species (e.g., surplus production, 

multi-species virtual population analyses, ecosystemlevel models such as ECOSIM and ECOPATH) should 

be evaluated. The natural mortality rate that is routinely 

applied in stock assessment models (18% a year) needs 

to be validated and the degree to which it varies for 

different age groups of herring, and in response to 

annual changes in the population abundance of herring, 

herring predators, and other prey species, should be 

determined. Also, stock assessment information could 

possibly be improved by developing a direct method for 

estimating annual changes in the abundance of juvenile 

herring that recruit to exploited stocks. 

More information is needed concerning the 

physical characteristics of benthic herring egg habitats 

and their vulnerability to disturbance by mobile, 

bottom-tending fishing gear, by natural disturbance, 

and, especially in nearshore spawning areas, to other 

habitat impacts related to human activities that are not 

associated with fishing. 

In the pelagic realm, more information is needed 

regarding oceanographic features that affect the 

abundance and distribution of larval, juvenile, and adult 

herring. Research in the northern North Sea (see 

Habitat Characteristics: Adults) has demonstrated that 

the abundance of pre-spawning adults is higher in 

oceanographic fronts between well-mixed and stratified 

water masses (such as exist along the edges of offshore 

banks) where the abundance of their zooplanktonic 

food supply is high. Given the relatively large amount 

of information that is available for this species, its 

importance as a prey species, and the fact that the 

juveniles and adults are amenable to acoustic survey 

procedures, the Atlantic herring is an excellent subject 

for pelagic habitat research. Most pelagic fisheriesrelated research in the Gulf of Maine region has been 

directed at factors affecting the distribution, growth, 

and survival of larvae, but not juvenile and adult fish. 

ACKNOWLEDGEMENTS 

The authors would like to thank Dave Packer and 

John McCarthy for editing, formatting, and graphics, 

and Jeff Pessuti, Barry Shafer, David Dowds, and Steve 

Fromm for mapping of the NEFSC groundfish data. 

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strategic issues for 4WX stock structure. In: 

Mooney-Seuss, M.L., Goebel, J.S., Tausig, H.C., 

Sweeney, M.S., editors. Herring stock assessment 

and research priorities. New England Aquarium 

Aquatic Forum Ser. Rep. 98-1. p. 8-19. 

Stephenson, R.L., K.J. Clark, M.J. Power, F.J. Fife, and 

G.D. Melvin. 2001. Herring stock structure, stock 

discreteness, and biodiversity. In: Funk, F., 

Blackburn, J., Hay, D., Paul, A.J., Stephenson, R., 

Toresen, R., Witherell, D., editors. Herring: 

expectations for a new millennium. Univ. Alaska 

Sea Grant , AK-SG-01-04, Fairbanks, AK. p. 559-

571. 

Stephenson, R.L., K. Rodman, D.G. Aldous, and D.E. 

Lane. 1999. An in-season approach to management 

under uncertainty: the case of the SW Nova Scotia 

herring fishery. ICES J. Mar. Sci. 56: 1005-1013. 

Stickney, A.P. 1972. The locomotor activity of juvenile 

herring (Clupea harengus harengus L.) in response 

to changes in illumination. Ecology 53:438-445. 

Sutcliffe, W.H., K. Drinkwater, and B.S. Muir. 1977. 

Correlations of fish catch and environmental 

factors in the Gulf of Maine. J. Fish. Res. Board 

Can. 34: 19-34. 

Tibbo, S.N. 1964. Effect of light on movements of 

herring in the Bay of Fundy. Intern. Comm. 

Northwest Atl. Fish. Spec. Pub. 6: 579-582. 

Wilson, K.W. 1974. The ability of herring and plaice 

larvae to avoid concentrations of oil dispersants. 

In: Blaxter, J.H.S., editor. The early life history of 

fish. Berlin: Springer-Verlag. p. 589-602. 

Winters, G.H. 1976. Recruitment mechanisms of 

southern Gulf of St. Lawrence Atlantic herring 

(Clupea harengus harengus). J. Fish. Res. Board 

Can. 33: 1751-1763. 

Zinkevich, V.N. 1967. Observations on the distribution 

of herring, Clupea harengus L., on Georges Bank 

and in adjacent waters in 1962-65. Intern. Comm. 

Northwest Atl. Fish. Res. Bull. 4: 101-115. 
Page 19

Table 1. Relative abundance of Atlantic herring life stages in New England and Mid-Atlantic estuaries and embayments 

by salinity zone. 

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   

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. 
Page 20

Table 1. Cont’d. 

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 

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. 
Page 21

Table 2. Distribution and abundance of juvenile Atlantic herring in New England and Mid-Atlantic estuaries and 

embayments. 

Month: 

Bay/River/Estuary: 

J F M A M J J A S O N D 

Passamaquoddy Bay A A A A H H H H H H A A 

Englishman/Machias Bays A A A A H H H H H A A A 

Narraguagus Bay A A A A H H H H H A A A 

Blue Hill Bay A A A A H H H H H A A A 

Penobscot Bay C C C A H H H H H H A C 

Muscongus Bay R A A A A A A C R 

Damariscotta River R A A A A C C C R 

Sheepscot River R A A A A C C C R 

Kennebec/Androscoggin Rivers R C C C C C C C R 

Casco Bay R A A A A A C C R 

Saco Bay R A A A A A C C R 

Wells Harbor C H H H H H A A R 

Great Bay C C C C C C C C R 

Merrimack River R C C C C C C C 

Massachusetts Bay A A A A A C C C A A A A 

Boston Harbor A A A A A C C C A A A A 

Cape Cod Bay A A A A A C C C C A A A 

Buzzards Bay C C C C C R R R R C C C 

Narragansett Bay C C C C C C C C C C C C 

Long Island Sound C C C C C C C C C C C C 

Gardiners Bay R R R R C C C C R R R R 

Great South Bay R R C C C C R R R R C C 

Hudson River/ Raritan Bay C C C C C R R R R R R R 

Barnegat Bay R R R C C C R R R R R R 

New Jersey Inland Bays R R R C C C R R R R R R 

Delaware Bay C C R R R R R 

Chincoteague Bay R R R 

Chesapeake Bay R R R 

Based on Estuarine Living Marine Resources (ELMR) data in Jury et al. (1994) and Stone et al. (1994). Relative 

abundance: H = highly abundant, A = abundant, C = common, R = rare, blank = not present. 
Page 22

Table 3. Distribution and abundance of adult Atlantic herring in New England and Mid-Atlantic estuaries and 

embayments. 

Month: 

Bay/River/Estuary: 

J F M A M J J A S O N D 

Passamaquoddy Bay A A A A H H H H H H A A 

Englishman/Machias Bays C C C A H H H H H A A C 

Narraguagus Bay C C C A H H H H H A A C 

Blue Hill Bay C C C A H H H H H A A C 

Penobscot Bay C C C A H H H H H H A C 

Muscongus Bay R C C C C A A C R 

Damariscotta River R C C C C A A C R 

Sheepscot River R C C C C A A C R 

Kennebec/Androscoggin Rivers R C C C C C C C R 

Casco Bay R C C C C C C C R 

Saco Bay R C C C C C C C R 

Wells Harbor R C C C C C C C R 

Great Bay R R R R R C C C R 

Merrimack River R R R R R R R 

Massachusetts Bay A A A A C R R R C C A A 

Boston Harbor A A A A A R R R C C C A 

Cape Cod Bay A A A A C R R R C C A A 

Buzzards Bay C C C C C R R R R C C C 

Narragansett Bay A A A A C C C C C C C C 

Long Island Sound A A A A A C C C C C A A 

Gardiners Bay R R R R C C C C R R R R 

Great South Bay A A C R R R R R R R A A 

Hudson River/ Raritan Bay C C C C C R R R R R R R 

Barnegat Bay C R C C 

New Jersey Inland Bays C R R R R R R R R R C C 

Delaware Bay C R R R R R R R R R C C 

Based on Estuarine Living Marine Resources (ELMR) data in Jury et al. (1994) and Stone et al. (1994). Relative 

abundance: H = highly abundant, A = abundant, C = common, R = rare, blank = not present. 
Page 23

Table 4. Percentage of Atlantic herring in the diets of 15 predators in the northeast U.S. Atlantic coast ecosystem. 

Percent herring 

in diet Taxon 

Predator species Size 

(cm) By wt By vol 

Years Location 

Number 

stomachs 

examined C. 

harengus Herrings Clupeidae 

Source 

15 1973-1975 

17 1976-1980 

2 1981-1985 

11 1986-1990 

51-120+ 

25 1991-1998 

8,176 

 over 

entire 

time 

period 

Link and Garrison 

(2002) 

61-70 4.4 86 

71-80 9.7 52 

Atlantic cod 

81-90 6.5 

1977-1980 

NE shelf 

91 

Bowman et al. (2000) 

< 20 4 8,722 

20-50 9 26,070 

> 50 25 

1973-1997 

1,037 

Garrison and Link 

(2000) 

26-30 4.0 323 

31-35 11.1 373 

41-45 20.5 72 

Silver hake 

> 45 23.3 

1977-1980 

NE shelf 

75 

Bowman et al. (2000) 

41-45 5.5 80 

56-60 13.4 

1977-1980 

44 Bowman et al. (2000) Summer flounder 

Mean = 36 8 1990-2000 

NE shelf 

na Link et al. (2002b) 

Atlantic halibut 41-50 11.1 1977-1980 26 Bowman et al. (2000) 

Mean = 58 4 1973-1998 

NE shelf 

155 Link et al. (2002b) 

51-60 2.5 235 

61-70 1.6 207 

71-80 8.3 697 

81-90 0.3 368 

Spiny dogfish 

91-100 1.3 

1977-1980 NE shelf 

423 

Bowman et al. (2000) 

20-50+ 20 1991-1997 na 

20-50 2 5,341 

White hake 

> 50 13 

1973-1997 

6,049 

Red hake > 50 2 1973-1997 

NE shelf 

1,713 

Garrison and Link 

(2000) 

Mean = 

221 87.2 Jeffreys Ledge 147 

Mean = 

221 48.4 Great South 

Channel 210 

Mean = 

240 6 Stellwagen Bank 111 

Mean = 

251 3.1 Cape Cod Bay 273 

Bluefin tuna 

Mean = 

124 2.5 

1988-1992 

South of 

Martha’s 

Vineyard 

57 

Chase (2002) 

11.3 1994 50 “Adults” 

17.6 1995 Georges Bank 44 Buckel et al. (1999) 

21-30 2.7 239 

Bluefish 

31-40 2.3 

1977-1980 NE shelf 

71 Bowman et al. (2000) 

30-120 3.4 North shore MA 1,536 

25-120 0.2 Cape Cod Bay 1,019 

Striped bass 

30-120 0 

1997-2000 

Nantucket Sound 451 

Nelson et al. (2003) 

Dusky shark 91-100 1.5 1977-1980 NE shelf 18 

61-70 36.5 36 

71-80 25.5 42 

Thorny skate 

> 90 20.8 

1977-1980 NE shelf 

18 

Bowman et al. (2000) 

51-60 1.9 104 

81-90 1.2 86 

Goosefish 

> 90 15.0 

1977-1980 NE shelf 

103 

Bowman et al. (2000) 

Black sea bass 21-25 2.3 1977-1980 NE shelf 188 Bowman et al. (2000) 

Weakfish 21-30 11.2 1977-1980 NE shelf 196 Bowman et al. (2000) 
Page 24

Table 5. Annual consumption (metric tons) of Atlantic herring by various predators. 

Fish and Elasmobranch Predators Marine Mammal Predators 

Species Estimated Annual Consumption, 

1977-1997 Species Estimated Annual Consumption, 1991-1997 

Spiny Dogfish 36,000-214,000 Fin Whale 16,081-62,362 

Silver Hake 11,500-36,000 Minke Whale 11,648-22,108 

Georges Bank Cod 1,900-13,000 Humpback Whale 31,046-35,507 

White Hake 500-20,000 Pilot Whale 149-512 

Bluefish 500-13,600 Harbor Porpoise 20,863-27,655 

Fluke 200-3,100 White-sided Dolphin 7,852-35,591 

Pollock 200-3,100 Harbor Seal 4,853 

Red Hake 200-3,100 Gray Seal 1,310 

Goosefish 200-3,100 

Winter Skate 200-3,100 

Gulf of Maine Cod 200-3,100 

 

Estimated Annual Consumption, 

1977-1998 

Spiny Dogfish 15,526-148,197 

(mean = 67,660) 

Winter Skate 20-2,329 

(mean = 928) 

Sources: Overholtz et al. (2000) (finfish and elasmobranchs, 1977-1997); Link et al. (2002a) (finfish and elasmobranchs, 

1977-1998); Read and Brownstein (2003) (marine mammals). 
Page 25

Table 6. Atlantic herring spawning site survey data and habitat parameters. 

Authors Observation or 

sampling 

method 

Location Date of 

survey 

Size of 

egg beds 

(km2

) 

Depth 

(m) 

McKenzie (1964) Biological 

dredge 

Trinity Ledge, 

southwest Nova Scotia 

Sept 1961 .067 11-13 

Caddy and Iles 

(1973) 

Submersible Northern edge of eastern 

Georges Bank 

Sept/Oct 

1970 

1.1 a

0.53 a

0.3 a

50 

Boyar et al. (1973) Dredge Jeffreys Ledge Oct 1972 53-59 

Cooper et al. 

(1975) 

SCUBA divers, 

grab samples 

Jeffreys Ledge Oct 1974 0.78 

1.39 

35-55 

Stevenson and 

Knowles (1988) 

ROV, grab 

samples, and 

SCUBA 

Eastern Maine Sept 1985, 

1986 

0.8 20-50 

Neal and Brehme 

(2001); Neal (2003) 

Small benthic 

sampler 

Eastern Maine 1997-2002 28-73 

Neal and Brehme 

(2001) 

Small benthic 

sampler 

Grand Manan Sept 2000 5-12 

P. Valentine, U.S. 

Geological Survey, 

Woods Hole Field 

Center, 384 Woods 

Hole Road, 

Quissett Campus, 

Woods Hole, MA 

02543, pers. comm. 

Underwater 

video, grab 

sample 

Stellwagen Bank Oct 1996 34 

a

 Egg bed sizes reported by Pankratov and Sigajev (1973). 

b

 Temperatures reported by Graham and Chenoweth (1973). 
Page 26

Table 6. Cont’d. 

Authors Substrate Bottom 

temp. 

(C) 

Bottom 

current 

(knots) 

Other 

McKenzie (1964) Flat, sandy bottom with few small 

stones, no vegetation; eggs also 

attached to an alga (Desmarestia 

aculeata) in a nearby site that was 

not surveyed. 

11.3-12 1.5-2 Egg mat 3.25 cm thick at deepest 

point, tapering to individual eggs 

spaced 2-6 mm apart; herring 

eggs in haddock stomachs. 

Caddy and Iles 

(1973)a

Pebbles 2-10 mm in diameter, 

boulders embedded in gravel, and 

on epifaunal growth; eggs thin or 

absent on sand. 

13-15b

 1-2 Eggs 1-2 cm thick in discrete bed. 

Predators (red hake, sculpin, 

dogfish, skate, hermit crabs, 

starfish, moon snails) left steep 

sided craters in egg layer – 8% 

eggs removed by predators. 

Boyar et al. 

(1973) 

Boulders, rocks, and gravel 7-8.5 Eggs collected in clumps, layers 

up to 5 mm thick. 

Cooper et al. 

(1975) 

80-90% eggs in Ptiloda serrata

attached to rocks, few on non-algal 

covered rock surfaces at one site. 

At another, bedrock, boulder, rock, 

gravel, shell, 10% Ptiloda on 

bedrock. 90% eggs on rock-gravel. 

Deeper than 55 m, fine sand, no 

eggs. 

9.5 0-1 Cunner most abundant predator 

observed feeding on eggs; red 

alga (Ptiloda) and eggs in cod 

stomachs. Hatching success > 

99% at one site where egg cover 

was sparse. 50-70% egg mortality 

at bottom of 4-5 cm thick egg 

mass. 

Stevenson and 

Knowles (1988) 

Egg cover thickest on gravel and 

shell fragments, very few or no 

eggs on rocks or fine sand/shells at 

edges of egg beds. 

 Eggs at two sites in continuous 

“carpet” 1-3 cm thick, in clumps 

and patches at two other sites. 

Egg mortality negligible, no signs 

of predation, egg development 

uniform throughout egg mass. 

 Neal and Brehme 

(2001); Neal 

(2003) 

Egg mats predominantly on gravel, 

eggs also observed on shell 

fragments and rocks 

 

P. Valentine, pers. 

comm. 

Coarse sand 
Page 27

Table 7. Summary of habitat characteristics and requirements for Atlantic herring in the northwest Atlantic continental 

shelf ecosystem. 

Eggs 

Habitat Discrete, demersal, egg “beds” in coastal waters and on offshore banks and ledges in the Gulf of Maine and 

on Georges Bank with strong bottom currents and coarse substrate. 

Depth 5-90 m. 

Substrate Boulders, rocks, gravel, coarse sand, shell fragments, macrophytes, and on a variety of benthic organisms 

and man-made structures (e.g., lobster traps); not on mud or fine sand. 

Temperature Bottom temperatures over egg beds ranged from 7-15oC; egg development normal 1-22oC; development 

rates/incubation times inversely related to temperature (10-15 days at 8-13oC). 

Salinity 32-33 ppt in situ, maximum hatching success 20-35 ppt (lab studies). 

Other Low hatching success at dissolved oxygen concentrations < 20% saturation; 100% mortality of eggs under 1 

cm of sediment, 85% mortality under thin film of sediment. 

Predators Cod, haddock, cunner, red hake, sand lance, probably moon snails, hermit crabs and starfish. 

Prey Not applicable. 

Larvae 

Habitat Pelagic, in estuaries, coastal, and offshore waters between Bay of Fundy and New Jersey; remain on or near 

bottom for first few days after hatching, until yolk-sac is absorbed, then rise to surface and are dispersed by 

currents. 

Depth Collected from very shallow water to 200 m, most 50-90 m. 

Substrate Not applicable. 

Temperature Lab study shows larvae tolerate wide temperature range (-1.8 to 24oC). 

Salinity Lab study shows larvae tolerate wide salinity range (2.5-52.5 ppt for 7 days). 

Other 50% mortality in dissolved oxygen concentrations of 1.9-3.6 mg/l at 10oC after 96 hours; crude oil in 

concentrations of 1-20 ml/l is toxic; narcotized by oil dispersants (lab studies). 

Predators Sand lance, jellyfish, Atlantic mackerel, Atlantic herring. 

Prey Developmental stages of copepods (7-20 mm larvae, in fall); small adult copepods (21-30 mm, in winter); 

wide variety of planktonic organisms (> 30 mm, in spring). 

Juveniles 

Habitat Pelagic; one-year-olds in nearshore waters during summer and fall, overwinter in deeper, coastal waters; twoyear-olds in inshore/offshore continental shelf waters of Gulf of Maine, deeper waters of Georges Bank in 

summer and fall, Cape Hatteras to deeper parts of Georges Bank in winter, widespread from Cape Hatteras to 

Bay of Fundy in spring. 

Depth Collected in bottom trawl surveys to edge of continental shelf (300 m), mostly < 100 m in spring; migrate up 

in water column at dusk and down at dawn. 

Substrate Not applicable. 

Temperature Lab studies show that juveniles prefer 8-12oC, physiological stress < 4oC and > 16o C, can survive -1.1oC, 

50% juveniles exposed to 19.5-21.2oC died within 48 hrs; most caught 3-7oC in spring and 6-10oC in fall 

NEFSC bottom trawl surveys. 

Salinity Lab studies show that juveniles prefer 28-32 ppt, can tolerate as low as 5 ppt for a short time, salinity 

preference is temperature-dependant (> 29 ppt below 10oC, no preference > 10oC); one-year-olds in 

coves/estuaries with low salinities, two-year-olds avoid brackish water. 

Other Spatial distribution affected by currents, frontal zones, and availability of zooplanktonic food organisms (see 

adults). 

Predators Heavily preyed upon by a variety of marine fish, marine mammals, and seabirds (see adults). 

Prey Feed on up to 15 types of zooplankton; most common are copepods, decapod larvae, barnacle larvae, 

cladocerans, and molluscan larvae. 
Page 28

Table 7. Cont’d. 

Adults 

Habitat Pelagic, but spawn on bottom; inshore/offshore continental shelf waters of the Gulf of Maine and deeper 

parts of Georges Bank in summer and fall, Cape Hatteras to deeper parts of Georges Bank in winter, 

distributed across shelf in mid-Atlantic, southern New England, deeper waters of Georges Bank, and the 

southwest portion of the Gulf of Maine in spring. 

Depth Collected in bottom trawl surveys to edge of continental shelf (300 m), mostly < 80 m in fall and at shallower 

depths in spring; diel vertical migrations similar to juveniles; more abundant < 150 m in northern North Sea 

in summer. 

Substrate Pre-spawning aggregations more abundant over gravel/sand in northern North Sea. 

Temperature Field observations suggest adults prefer 5-9oC on Georges Bank in summer/fall; most caught 4-7oC in spring 

and 6-10oC in fall NEFSC trawl surveys; adults more abundant in areas of northern North Sea where summer 

sea surface temperatures are 11-14oC, thermocline 25-45 m deep, and difference between surface and bottom 

water temperatures was only 3oC. 

Salinity Adults most abundant 27-35 ppt in spring NEFSC trawl surveys, 32-34 ppt at other times of year. 

Other Well-mixed (unstratified) waters and transition zones (fronts) between well-mixed and stratified waters are 

preferred habitats for adults; also more abundant in or on edges of plankton “patches.” 

Predators Important forage species in NW Atlantic ecosystem; principal finfish and elasmobranch predators are cod, 

silver hake, thorny skate, bluefish, monkfish, weakfish, summer flounder, white hake, and spiny dogfish – 

also, at certain locations and times of year – Atlantic bluefin tuna; principal marine mammal predators are 

minke whales, harbor porpoise, white-sided dolphins, fin and humpback whales. 

Prey Principal zooplankton prey organisms are euphausiids, amphipods, copepods, chateognaths, pteropods, 

mysids, and pandalid shrimp; adults also consume fish eggs and larvae (including their own). 
Page 29

Figure 1. The Atlantic herring, Clupea harengus L. (from Goode 1884). 
Page 30

#

Cape Hatteras

36°

36°

38°

38°

40°

40°

42°

42°

44°

44°

76°

76°

74°

74°

72°

72°

70°

70°

68°

68°

66°

66°

Outer Limit

 of EEZ

100 m

Gulf of Canada

 Maine

Georges 

 Bank

Mid-Atlantic

 Bight

#

Cape Cod

#

Long Island

#

Delaware Bay

#

Chesapeake Bay

200 0 200 400 Miles

#

Cape Hatteras

Figure 2. Northeast U.S. Atlantic coast ecosystem. 
Page 31

Figure 3. Generalized view of principal Atlantic herring spawning areas in the Gulf of Maine and on Georges Bank. 

Source: Overholtz et al. 2004. 

-71 -70 -69 -68 -67 -66 -65 -64

40

41

42

43

44

45

 

 

Nantucket

Shoals

Georges

Bank

Bay of

Fundy Grand

Manan

Jeffreys

Ledge

Western Maine

Penobscot

Bay

Scots Bay

Eastern Maine
Page 32

Figure 4. Percent by weight of the major prey items in the diet of Atlantic herring. Specimens were collected during 

NEFSC bottom trawl surveys from 1973-2001 (all seasons). For details on NEFSC diet analysis, see Link and Almeida 

(2000). 

0

10

20

30

40

Hyperid

Amphipods

Well

Digested

Prey

Chaetognaths

Copepods

Misc.

Crustaceans

Decapods

Euphasiids

Larvaceans

Mysids

Percent
Page 33

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

Sindermann (1979). 
Page 34

Figure 6. Geo-referenced in situ observations of Atlantic herring eggs (see Table 6). 

#

# # #

### #

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#

#### #

###

###

#

Cape Cod

Maine

MA

NH

#

Penobscot 

 Bay #

Casco

 Bay

GULF OF

 MAINE

US

CAN

Bay of

Fundy

#

Stellwagen

 Bank

#

Jeffreys

 Ledge

Georges Bank

Nantucket 

 Shoals

40°

40°

41°

41°

42°

42°

43°

43°

44°

44°

45°

45°

71°

71°

70°

70°

69°

69°

68°

68°

67°

67°

66°

66°

 Nova

Scotia

100 m
Page 35

Figure 7. Atlantic herring spawning sites in eastern Maine, 1997-2002. Depth contours in 10 m intervals. Source: 

Island Institute, Rockland, ME [see Neal and Brehme (2001) and Neal (2003)]. 

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Machias Bay

#

Cutler

MAINE

70m

#

Cross

Island

#

 West

Quoddy

 Head
Page 36

Figure 8. Principal spawning grounds of Atlantic herring on Georges Bank, 1964-1971 (excluding 1967), with a 

comparison of egg patch sizes among years. Source: Anthony and Waring (1980). 
Page 37

(Clupea harengus)

MARMAP Ichthyoplankton Surveys

61-cm Bongo Net; 0.505-mm mesh

January - December, 1977 to 1987

Atlantic Herring

Number of Tows = 11438, with larvae = 607

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

Figure 9. Distributions and abundances of Atlantic herring larvae collected during NEFSC MARMAP ichthyoplankton 

surveys. 

For all available months and years from 1977 to 1987 combined. 
Page 38

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

35

36

37

38

39

40

41

42

43

44

45

January, 1977 to 1987

Number of Tows = 434, with larvae = 39

Monthly Mean Density = 2.51 Larvae/10m2

Number of Larvae / 10m2

None

1 to < 10

10 to < 100

100 to 174

February, 1977 to 1987

Number of Tows = 686, with larvae = 62

Monthly Mean Density = 0.68 Larvae/10m2

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

March, 1977 to 1987

Number of Tows = 1031, with larvae = 14

Monthly Mean Density = 0.11 Larvae/10m2

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

April, 1977 to 1987

Number of Tows = 1281, with larvae = 28

Monthly Mean Density = 0.21 Larvae/10m2

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

Figure 9. Cont’d. 

From MARMAP ichthyoplankton surveys, January through April, 1977-1987. 
Page 39

May, 1977 to 1987

Number of Tows = 1472, with larvae = 3

Monthly Mean Density = 0.01 Larvae/10m2

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

June, 1977 to 1987

Number of Tows =893, with larvae = 2

Monthly Mean Density = 0.01 Larvae/10m2

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

July, 1977 to 1987

Number of Tows =938, with larvae = 1

Monthly Mean Density = 0.01 Larvae/10m2

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

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

35

36

37

38

39

40

41

42

43

44

45

August, 1977 to 1987

Number of Tows = 1148, with larvae = 2

Monthly Mean Density = 0.07 Larvae/10m2

Number of Larvae / 10m2

None

1 to < 10

10 to 70

Figure 9. Cont’d. 

From MARMAP ichthyoplankton surveys, May through August, 1977-1987. 
Page 40

September, 1977 to 1987

Number of Tows = 774, with larvae = 28

Monthly Mean Density = 350.82 Larvae/10m2

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

October, 1977 to 1987

Number of Tows = 1147, with larvae = 117

Monthly Mean Density = 39.30 Larvae/10m2

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

November, 1977 to 1987

Number of Tows = 1031, with larvae = 202

Monthly Mean Density = 49.07 Larvae/10m2

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

December, 1977 to 1987

Number of Tows = 603, with larvae = 109

Monthly Mean Density = 5.41 Larvae/10m2

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

Figure 9. Cont’d. 

From MARMAP ichthyoplankton surveys, September through December, 1977-1987. 
Page 41

Figure 10. Changes in abundance of Atlantic herring larvae on Georges Bank, Nantucket Shoals, and in Massachusetts 

Bay from 1971-1990. 

Source: Smith and Morse (1993). Intervals (Int.) denote periods of changing spawning patterns. 
Page 42

Figure 11. Distribution of Atlantic herring larvae by age in the Georges Bank area, 1971-1990. 

Source: Smith and Morse (1993). 
Page 43

Figure 11. Cont’d. 
Page 44

Figure 12. Distribution of recently-hatched Atlantic herring larvae on Georges Bank, 1988-1994. 

Source: Melvin et al. (1996). 
Page 45

Figure 13. Seasonal distributions and abundances of juvenile Atlantic herring collected during NEFSC bottom trawl 

surveys. 

From NEFSC summer bottom trawl surveys (1963-1995, all years combined). Distributions are displayed as 

presence/absence only. 
Page 46

Figure 13. Cont’d. 

From NEFSC fall bottom trawl surveys (1963-2003, all years combined). Survey stations where juveniles were not 

found are not shown. 
Page 47

Figure 13. Cont’d. 

From NEFSC winter bottom trawl surveys (1964-2003, all years combined). Distributions are displayed as 

presence/absence only. 
Page 48

Figure 13. Cont’d. 

From NEFSC spring bottom trawl surveys (1968-2003, all years combined). Survey stations where juveniles were not 

found are not shown. 
Page 49

Figure 14. Seasonal distributions and abundances of adult Atlantic herring collected during NEFSC bottom trawl 

surveys. 

From NEFSC summer bottom trawl surveys (1963-1995, all years combined). Distributions are displayed as 

presence/absence only. 
Page 50

Figure 14. Cont’d. 

From NEFSC fall bottom trawl surveys (1963-2003, all years combined). Survey stations where adults were not found 

are not shown. 
Page 51

Figure 14. Cont’d. 

From NEFSC winter bottom trawl surveys (1964-2003, all years combined). Distributions are displayed as 

presence/absence only. 
Page 52

Figure 14. Cont’d. 

From NEFSC spring bottom trawl surveys (1968-2003, all years combined). Survey stations where adults were not found 

are not shown. 
Page 53

Figure 15. Distribution and abundance of Atlantic herring along the coasts of Maine and New Hampshire during spring 

of 2001-2003 and fall 2000-2002, from the Maine – New Hampshire inshore groundfish trawl survey. For details on the 

survey, see Sherman et al. (2004). 
Page 54

Figure 16. Length frequency plots for Atlantic herring caught along the Maine and New Hampshire coasts, by 

season/year. Based on the Maine – New Hampshire inshore groundfish trawl survey for spring 2001-2003 and fall 2000-

2002. Source: Sherman et al. (2004). 
Page 55

Figure 17. Seasonal distributions and abundances of juvenile Atlantic herring in Massachusetts coastal waters. 

From spring Massachusetts inshore bottom trawl surveys (1978-2003, all years combined). Survey stations where 

juveniles were not found are not shown. 
Page 56

Figure 17. Cont’d. 

From fall Massachusetts inshore bottom trawl surveys (1978-2003, all years combined). Survey stations where juveniles 

were not found are not shown. 
Page 57

Figure 18. Seasonal distributions and abundances of adult Atlantic herring in Massachusetts coastal waters. 

From spring Massachusetts inshore bottom trawl surveys (1978-2003, all years combined). Survey stations where adults 

were not found are not shown. 
Page 58

Figure 18. Cont’d. 

From fall Massachusetts inshore bottom trawl surveys (1978-2003, all years combined). Survey stations where adults 

were not found are not shown. 
Page 59

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)

Figure 19. Seasonal distribution and abundance of juvenile Atlantic herring in Narragansett Bay. 

Based upon the Rhode Island bottom trawl surveys, 1990-1996. The numbers shown at each station are the average 

catch per tow rounded to one decimal place [see Reid et al. (1999b) for details]. 
Page 60

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)

Figure 20. Seasonal distribution and abundance of adult Atlantic herring in Narragansett Bay. 

Based upon the Rhode Island bottom trawl surveys, 1990-1996. The numbers shown at each station are the average 

catch per tow rounded to one decimal place [see Reid et al. (1999b) for details]. 
Page 61

Figure 21. Distribution and abundances of juvenile and adult Atlantic herring in Long Island Sound. 

Based on the finfish surveys of the Connecticut Fisheries Division, 1984-1994 [from Gottschall et al. (2000)]. Circle 

diameter is proportional to the number of fish caught, and is scaled to the maximum catch (indicated by “max=” or 

“max>”). Collections were made with a 14 m otter trawl at about 40 stations chosen by stratified random design. 
Page 62

Figure 22. Relative abundance (geometric mean catch/tow) catch/tow and percent occurrence (proportion of samples in 

which at least one individual was observed) for juvenile and adult Atlantic herring in Long Island Sound. 

By month, month and bottom type, and month and depth interval. From Gottschall et al. (2000). 
Page 63

Figure 23. Monthly log10 length frequencies (cm) of juvenile and adult Atlantic herring collected in Long Island Sound. 

Based on 21,149 fish taken in 360 tows between 1989 and 1994. From Gottschall et al. (2000). 
Page 64

Figure 24. Seasonal distribution and abundance of juvenile Atlantic herring in the Hudson-Raritan Estuary. 

Based on Hudson-Raritan trawl surveys, January 1992 – June 1997 [see Reid et al. (1999b) for details]. 
Page 65

Figure 25. Seasonal distribution and abundance of adult Atlantic herring in the Hudson-Raritan Estuary. 

Based on Hudson-Raritan trawl surveys, January 1992 – June 1997 [see Reid et al. (1999b) for details]. 
Page 66

 

Figure 26. Catch per unit effort for total catch of juvenile and adult Atlantic herring in Chesapeake Bay, from the 

Virginia Institute of Marine Science’s (VIMS) trawl surveys, 1988-1999 (all years combined). Monthly surveys were 

conducted using a random stratified design of the main stem of the Bay using a 9.1 m semi-balloon otter trawl with 38 

mm mesh and 6.4 mm cod end with a tow duration of five minutes. Source: Geer (2002). 
Page 67

Figure 27. Seasonal distribution and abundance of juvenile Atlantic herring in Chesapeake Bay, from the VIMS trawl 

surveys, 1988-1999 (all years combined). Source: Geer (2002). 
Page 68

Figure 28. Seasonal distribution and abundance of adult Atlantic herring in Chesapeake Bay, from the VIMS trawl 

surveys, 1988-1999 (all years combined). Source: Geer (2002). 
Page 69

Figure 29. Atlantic herring catch per unit effort by site from the VIMS beach seine surveys, 1994-1999 (all years 

combined). Source: Geer (2002).
Page 70

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

1030

507090

110

130150

170

190

210

230250270290

325375

450750

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

1030507090

110130

150

170190

210

230

250

270

290325375

450750

1250

1750

>2000

0

10

20

30

90

100

Figure 30. Distributions of Atlantic herring larvae collected during NEFSC MARMAP icthyoplankton surveys relative 

to water column temperature and bottom depth. 

For the years 1977-1987, by month for all years combined. Open bars represent the proportion of all stations which were 

surveyed, while solid bars represent the proportion of the sum of all standardized catches (number/10 m2

). Note that the 

bottom depth interval changes with increasing depth. 
Page 71

Atlantic Herring

NEFSC Bottom Trawl Survey

Spring 1968 - 2003

Juveniles (<25 cm)

0

2

4

6

8

10

12

14

16

1-10

11-2021-3031-4041-5051-6061-7071-8081-90

91-100

101-120121-140141-160161-180181-200201-300301-400401-500

>500

Bottom Depth (m)

Percent

Trawls N=12514

Occurrence N=3444

Catch N=184284

0

10

20

30

40

50

60

24 25 26 27 28 29 30 31 32 33 34 35 36 37 38

Salinity (PPT)

Percent

Trawls N=2270

Occurrence N=1051

Catch N=83471

0

5

10

15

20

25

30

35

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30

Bottom Temperature (°C)

Percent

Trawls N=10879

Occurrence N=3026

Catch N=168096

Figure 31. Distributions of juvenile Atlantic herring and trawls from NEFSC bottom trawl surveys relative to bottom 

water temperature, depth, and salinity. 

Based on NEFSC spring bottom trawl surveys (temperature and depth: 1968-2003, all years combined; salinity: 1991-

2003, all years combined). Light bars show the distribution of all the trawls, dark bars show the distribution of all trawls 

in which Atlantic herring occurred and medium bars show, within each interval, the percentage of the total number of 

Atlantic herring caught. Note that the bottom depth interval changes with increasing depth. 
Page 72

Atlantic Herring

NEFSC Bottom Trawl Survey

Fall 1963 - 2003

Juveniles (<25 cm)

0

5

10

15

20

25

1-10

11-2021-3031-4041-5051-6061-7071-8081-90

91-100

101-120121-140141-160161-180181-200201-300301-400401-500

>500

Bottom Depth (m)

Percent

Trawls N=14186

Occurrence N=1364

Catch N=78453

0

10

20

30

40

50

60

70

24 25 26 27 28 29 30 31 32 33 34 35 36 37 38

Salinity (PPT)

Percent

Trawls N=2272

Occurrence N=506

Catch N=23087

0

10

20

30

40

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30

Bottom Temperature (°C)

Percent

Trawls N=12207

Occurrence N=1145

Catch N=55849

Figure 31. Cont’d. 

Based on NEFSC fall bottom trawl surveys (temperature and depth: 1963-2003, all years combined; salinity: 1991-2003, 

all years combined). Light bars show the distribution of all the trawls, dark bars show the distribution of all trawls in 

which Atlantic herring occurred and medium bars show, within each interval, the percentage of the total number of 

Atlantic herring caught. Note that the bottom depth interval changes with increasing depth. 
Page 73

Atlantic Herring

Massachusetts Inshore Trawl Survey

Spring 1978 - 2003

Juveniles (<25 cm)

0

5

10

15

20

25

30

35

40

1-5

6-10

11-1516-2021-2526-3031-3536-4041-4546-5051-5556-6061-6566-7071-7576-8081-85

Bottom Depth (m)

Percent

Trawls N=2482

Occurrence N=481

Catch N=79413

0

5

10

15

20

25

30

35

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30

Bottom Temperature (°C)

Percent

Trawls N=2407

Occurrence N=470

Catch N=79269

Figure 32. Distributions of juvenile Atlantic herring and trawls in Massachusetts coastal waters relative to bottom water 

temperature and depth. 

Based on spring Massachusetts inshore bottom trawl surveys (1978-2003, all years combined). Light bars show the 

distribution of all the trawls, dark bars show the distribution of all trawls in which Atlantic herring occurred and medium 

bars show, within each interval, the percentage of the total number of Atlantic herring caught. 
Page 74

Atlantic Herring

Massachusetts Inshore Trawl Survey

Fall 1978 - 2003

Juveniles (<25 cm)

0

10

20

30

40

50

60

70

80

1-5

6-10

11-1516-2021-2526-3031-3536-4041-4546-5051-5556-6061-6566-7071-7576-8081-85

Bottom Depth (m)

Percent

Trawls N=2338

Occurrence N=376

Catch N=29742

0

10

20

30

40

50

60

70

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30

Bottom Temperature (°C)

Percent

Trawls N=2244

Occurrence N=362

Catch N=29128

Figure 32. Cont’d. 

Based on fall Massachusetts inshore bottom trawl surveys (1978-2003, all years combined). Light bars show the 

distribution of all the trawls, dark bars show the distribution of all trawls in which Atlantic herring occurred and medium 

bars show, within each interval, the percentage of the total number of Atlantic herring caught. 
Page 75

Figure 33. Distributions of juvenile Atlantic herring in Narragansett Bay relative to mean bottom temperature and 

bottom depth. 

Based on the Rhode Island bottom trawl survey, 1990-1996. Open bars represent stations surveyed and closed bars 

represent fish collected. 
Page 76

Figure 34. Distributions of juvenile Atlantic herring in the Hudson-Raritan Estuary relative to mean water temperature, 

depth, dissolved oxygen, and salinity. 

Based on the Hudson-Raritan trawl surveys, 1992-1997. Open bars represent stations surveyed and closed bars represent 

fish collected. 
Page 77

Figure 35. Hydrographic preferences for juvenile Atlantic herring in Chesapeake Bay, from the VIMS trawl surveys, 

1988-1999 (all years combined). Source: Geer (2002).
Page 78

Atlantic Herring

NEFSC Bottom Trawl Survey

Spring 1968 - 2003

Adults (>=25 cm)

0

5

10

15

20

25

1-10

11-2021-3031-4041-5051-6061-7071-8081-90

91-100

101-120121-140141-160161-180181-200201-300301-400401-500

>500

Bottom Depth (m)

Percent

Trawls N=12514

Occurrence N=3227

Catch N=84332

0

10

20

30

40

50

60

24 25 26 27 28 29 30 31 32 33 34 35 36 37 38

Salinity (PPT)

Percent

Trawls N=2270

Occurrence N=918

Catch N=19663

0

5

10

15

20

25

30

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30

Bottom Temperature (°C)

Percent

Trawls N=10879

Occurrence N=2825

Catch N=72071

Figure 36. Distributions of adult Atlantic herring and trawls from NEFSC bottom trawl surveys relative to bottom water 

temperature, depth, and salinity. 

Based on NEFSC spring bottom trawl surveys (temperature and depth: 1968-2003, all years combined; salinity: 1991-

2003, all years combined). Light bars show the distribution of all the trawls, dark bars show the distribution of all trawls 

in which Atlantic herring occurred and medium bars show, within each interval, the percentage of the total number of 

Atlantic herring caught. Note that the bottom depth interval changes with increasing depth. 
Page 79

Atlantic Herring

NEFSC Bottom Trawl Survey

Fall 1963 - 2003

Adults (>=25 cm)

0

5

10

15

20

1-10

11-2021-3031-4041-5051-6061-7071-8081-90

91-100

101-120121-140141-160161-180181-200201-300301-400401-500

>500

Bottom Depth (m)

Percent

Trawls N=14186

Occurrence N=1639

Catch N=74283

0

10

20

30

40

50

60

70

24 25 26 27 28 29 30 31 32 33 34 35 36 37 38

Salinity (PPT)

Percent

Trawls N=2272

Occurrence N=486

Catch N=26889

0

5

10

15

20

25

30

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30

Bottom Temperature (°C)

Percent

Trawls N=12207

Occurrence N=1401

Catch N=59811

Figure 36. Cont’d. 

Based on NEFSC fall bottom trawl surveys (temperature and depth: 1963-2003, all years combined; salinity: 1991-2003, 

all years combined). Light bars show the distribution of all the trawls, dark bars show the distribution of all trawls in 

which Atlantic herring occurred and medium bars show, within each interval, the percentage of the total number of 

Atlantic herring caught. Note that the bottom depth interval changes with increasing depth. 
Page 80

Atlantic Herring

Massachusetts Inshore Trawl Survey

Spring 1978 - 2003

Adults (>=25 cm)

0

5

10

15

20

25

30

1-5

6-10

11-1516-2021-2526-3031-3536-4041-4546-5051-5556-6061-6566-7071-7576-8081-85

Bottom Depth (m)

Percent

Trawls N=2482

Occurrence N=110

Catch N=368

0

10

20

30

40

50

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30

Bottom Temperature (°C)

Percent

Trawls N=2407

Occurrence N=106

Catch N=351

Figure 37. Distributions of adult Atlantic herring and trawls in Massachusetts coastal waters relative to bottom water 

temperature and depth. 

Based on spring Massachusetts inshore bottom trawl surveys (1978-2003, all years combined). Light bars show the 

distribution of all the trawls, dark bars show the distribution of all trawls in which Atlantic herring occurred and medium 

bars show, within each interval, the percentage of the total number of Atlantic herring caught. 
Page 81

Atlantic Herring

Massachusetts Inshore Trawl Survey

Fall 1978 - 2003

Adults (>=25 cm)

0

5

10

15

20

25

30

35

40

1-5

6-10

11-1516-2021-2526-3031-3536-4041-4546-5051-5556-6061-6566-7071-7576-8081-85

Bottom Depth (m)

Percent

Trawls N=2338

Occurrence N=80

Catch N=785

0

10

20

30

40

50

60

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30

Bottom Temperature (°C)

Percent

Trawls N=2244

Occurrence N=79

Catch N=777

Figure 37. Cont’d. 

Based on fall Massachusetts inshore bottom trawl surveys (1978-2003, all years combined). Light bars show the 

distribution of all the trawls, dark bars show the distribution of all trawls in which Atlantic herring occurred and medium 

bars show, within each interval, the percentage of the total number of Atlantic herring caught. 
Page 82

Figure 38. Distributions of adult Atlantic herring in Narragansett Bay relative to mean bottom temperature and bottom 

depth. 

Based on the Rhode Island bottom trawl survey, 1990-1996. Open bars represent stations surveyed and closed bars 

represent fish collected. 
Page 83

Figure 39. Distributions of adult Atlantic herring in the Hudson-Raritan Estuary relative to mean water temperature, 

depth, dissolved oxygen, and salinity. 

Based on the Hudson-Raritan trawl surveys, 1992-1997. Open bars represent stations surveyed and closed bars represent 

fish collected. 
Page 84

Figure 40. Hydrographic preferences for adult Atlantic herring in Chesapeake Bay, from the VIMS trawl surveys, 1988-

1999 (all years combined). Source: Geer (2002). 
Publishing in NOAA Technical Memorandum NMFS-NE

Manuscript Qualification

This series represents a secondary level of scientific

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For all issues, the series employs thorough internal scientific

review, but not necessarily external scientific review. For most

issues, the series employs rigorous technical and copy editing.

Manuscripts that may warrant a primary level of scientific

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Identical, or fundamentally identical, manuscripts should

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Manuscripts by Northeast Fisheries Science Center

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For spelling of scientific and common names of fishes, mollusks, and decapod crustaceans from the United States and Canada,

use Special Publications No. 29 (fishes), 26 (mollusks), and 17

(decapod crustaceans) of the American Fisheries Society (Bethesda,

MD). For spelling of scientific and common names of marine

mammals, use Special Publication No. 4 of the Society for Marine

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Publications and Reports

of the

Northeast Fisheries Science Center

The mission of NOAA's National Marine Fisheries Service (NMFS) is "stewardship of living marine resources

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NOAA Technical Memorandum NMFS-NE -- This series is issued irregularly. The series typically includes: data reports of longterm field or lab studies of important species or habitats; synthesis reports for important species or habitats; annual reports of overall

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MEDIA

 MAIL