NOAA Technical Memorandum NMFS-NE-145

Essential Fish Habitat Source Document:

Butterfish, Peprilus triacanthus,

Life History and Habitat Characteristics

U. S. DEPARTMENT OF COMMERCE

National Oceanic and Atmospheric Administration

National Marine Fisheries Service

Northeast Region

Northeast Fisheries Science Center

Woods Hole, Massachusetts

September 1999
Recent Issues

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Access. No. PB98-119159.

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

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

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

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

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

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

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

Access. No. PB98-112345.

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

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

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

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

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

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

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

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

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

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

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

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

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

William Daley, Secretary

National Oceanic and Atmospheric Administration

D. James Baker, Administrator

National Marine Fisheries Service

Penelope D. Dalton, Assistant Administrator for Fisheries

Northeast Region

Northeast Fisheries Science Center

Woods Hole, Massachusetts

September 1999

Essential Fish Habitat Source Document:

Butterfish, Peprilus triacanthus,

Life History and Habitat Characteristics

Jeffrey N. Cross1, Christine A. Zetlin1, Peter L. Berrien1,

Donna L. Johnson1, and Cathy McBride2

1

National Marine Fisheries Serv., James J. Howard Marine Sciences Lab., 74 Magruder Rd., Highlands, NJ 07732 2

Inst. for Marine and Coastal Studies, Rutgers Univ., Cook Campus, New Brunswick, NJ 08903

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

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

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

All issues may be cited in formal scientific communications.

NOAA Technical Memorandum NMFS-NE-145
Editorial Notes on Issues 122-152

in the

NOAA Technical Memorandum NMFS-NE Series

Editorial Production

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

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

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

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

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

Special Acknowledgments

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

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

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

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

152.

Internet Availability

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

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

Information Updating

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

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

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

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

Species Names

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

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

Turgeon et al. 1998b

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

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

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

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

).

a

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

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

b

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

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

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

c

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

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

d

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

e

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

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

FOREWORD

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

commercial and recreational fisheries is the continuing

loss of marine, estuarine, and other aquatic habitats.

Magnuson-Stevens Fishery Conservation and

Management Act (October 11, 1996)

The long-term viability of living marine resources

depends on protection of their habitat.

NMFS Strategic Plan for Fisheries

Research (February 1998)

The Magnuson-Stevens Fishery Conservation and

Management Act (MSFCMA), which was reauthorized

and amended by the Sustainable Fisheries Act (1996),

requires the eight regional fishery management councils to

describe and identify essential fish habitat (EFH) in their

respective regions, to specify actions to conserve and

enhance that EFH, and to minimize the adverse effects of

fishing on EFH. Congress defined EFH as “those waters

and substrate necessary to fish for spawning, breeding,

feeding or growth to maturity.” The MSFCMA requires

NMFS to assist the regional fishery management councils

in the implementation of EFH in their respective fishery

management plans.

NMFS has taken a broad view of habitat as the area

used by fish throughout their life cycle. Fish use habitat

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

most habitats provide only a subset of these functions.

Fish may change habitats with changes in life history

stage, seasonal and geographic distributions, abundance,

and interactions with other species. The type of habitat,

as well as its attributes and functions, are important for

sustaining the production of managed species.

The Northeast Fisheries Science Center compiled the

available information on the distribution, abundance, and

habitat requirements for each of the species managed by

the New England and Mid-Atlantic Fishery Management

Councils. That information is presented in this series of

30 EFH species reports (plus one consolidated methods

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

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

HIGHLANDS, NEW JERSEY

SEPTEMBER 1999

independent data sets from NMFS and several coastal

states. The species reports are also the source for the

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

understandably begun to be referred to as the “EFH source

documents.”

NMFS provided guidance to the regional fishery

management councils for identifying and describing EFH

of their managed species. Consistent with this guidance,

the species reports present information on current and

historic stock sizes, geographic range, and the period and

location of major life history stages. The habitats of

managed species are described by the physical, chemical,

and biological components of the ecosystem where the

species occur. Information on the habitat requirements is

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

available, habitat and environmental variables that control

or limit distribution, abundance, growth, reproduction,

mortality, and productivity.

Identifying and describing EFH are the first steps in

the process of protecting, conserving, and enhancing

essential habitats of the managed species. Ultimately,

NMFS, the regional fishery management councils, fishing

participants, Federal and state agencies, and other

organizations will have to cooperate to achieve the habitat

goals established by the MSFCMA.

A historical note: the EFH species reports effectively

recommence a series of reports published by the NMFS

Sandy Hook (New Jersey) Laboratory (now formally

known as the James J. Howard Marine Sciences

Laboratory) from 1977 to 1982. These reports, which

were formally labeled as Sandy Hook Laboratory

Technical Series Reports, but informally known as “Sandy

Hook Bluebooks,” summarized biological and fisheries

data for 18 economically important species. The fact that

the bluebooks continue to be used two decades after their

publication persuaded us to make their successors – the 30

EFH source documents – available to the public through

publication in the NOAA Technical Memorandum NMFSNE series.

JEFFREY N. CROSS, CHIEF

ECOSYSTEMS PROCESSES DIVISION

NORTHEAST FISHERIES SCIENCE CENTER
Page v

Contents

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

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

Habitat Characteristics ..............................................................................................................................................................................3

Geographical Distribution .........................................................................................................................................................................4

Status of the Stocks ...................................................................................................................................................................................6

Acknowledgments.....................................................................................................................................................................................6

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

Tables

Table 1. Summary of life history and habitat characteristics for butterfish, Peprilus triacanthus.......................................................11

Table 2a. Relative abundance of butterfish eggs, larvae, and juveniles in New England/Mid-Atlantic estuaries by salinity zone .......13

Table 2b. Relative abundance of butterfish adults and spawning adults in New England/Mid-Atlantic estuaries by salinity zone ......15

Table 3. Abundance of butterfish in New England and Mid-Atlantic estuaries by month summarized across salinity zones.............17

Figures

Figure 1. The adult butterfish, Peprilus triacanthus (from Goode 1884) .............................................................................................18

Figure 2. The overall distribution of butterfish from Newfoundland to Cape Hatteras.........................................................................19

Figure 3. Abundance (% occurrence) of the major prey items of butterfish collected during NEFSC bottom trawl surveys ...............20

Figure 4. Water temperature and depth where butterfish eggs were collected in the NEFSC MARMAP ichthyoplankton surveys.....21

Figure 5. Water temperature and depth where butterfish larvae were collected in the NEFSC MARMAP ichthyoplankton surveys ..22

Figure 6. Water temperature and depth where juvenile and adult butterfish were collected in the NEFSC bottom trawl surveys .......23

Figure 7. Water temperature and depth where juvenile and adult butterfish were collected in the Massachusetts trawl surveys .........25

Figure 8. Water temperature, depth, DO, and salinity where juvenile and adult butterfish were collected in the Hudson-Raritan ......26

Figure 9. Distribution and abundance of butterfish eggs based on NEFSC MARMAP ichthyoplankton surveys................................27

Figure 10. Distribution and abundance of butterfish larvae collected during NEFSC MARMAP ichthyoplankton surveys..................29

Figure 11. Distribution and abundance of juvenile and adult butterfish collected during NEFSC bottom trawl surveys .......................32

Figure 12. Distribution of juvenile and adult butterfish in Massachusetts coastal waters during Massachusetts trawl surveys..............34

Figure 13. Distribution of juvenile and adult butterfish in the SEAMAP bottom trawl surveys.............................................................35

Figure 14. Distribution of juvenile and adult butterfish collected in Narragansett Bay during Rhode Island bottom trawl surveys.......37

Figure 15. Distribution of juvenile and adult butterfish in Long Island Sound, from the Connecticut bottom trawl surveys.................39

Figure 16. Distribution of juvenile and adult butterfish in the Hudson-Raritan estuary based on Hudson-Raritan trawl surveys. .........40

Figure 17. Commercial landings and NEFSC bottom trawl indices for butterfish from the Gulf of Maine to the Middle Atlantic........42
Page 1

INTRODUCTION

Butterfish, Peprilus triacanthus (Figure 1), range

from Newfoundland and the Gulf of St. Lawrence to the

Atlantic and Gulf coasts of Florida (Figure 2), but they

are most abundant from the Gulf of Maine to Cape

Hatteras (Bigelow and Schroeder 1953; Haedrich 1967;

Horn 1970a; Powell et al. 1972; Cooley 1978; Scott and

Scott 1988; Brodziak 1995; Klein-MacPhee, in review).

Butterfish are fast-growing, short-lived, pelagic fishes that

form loose schools, often near the surface (Schreiber

1973; Dery 1988; Brodziak 1995). They winter near the

edge of the continental shelf in the Middle Atlantic Bight

and migrate inshore in the spring into southern New

England and Gulf of Maine waters. During the summer,

butterfish occur over the entire mid-Atlantic shelf from

sheltered bays and estuaries out to about 200 m. In late

fall, butterfish move southward and offshore in response

to falling water temperatures (Fritz 1965; Horn 1970a;

Schreiber 1973; Waring 1975; Azarovitz et al. 1980;

Klein-MacPhee, in review).

LIFE HISTORY

Butterfish are short-lived and grow rapidly; few

individuals live beyond 3 years and most are sexually

mature at 1-2 years of age. The maximum age reported is

3+ years (DuPaul and McEachran 1973; Waring 1975;

Kawahara 1977a) and 6 years (Draganik and Zukowski

1966). Butterfish are eurythermal (4.4-21.6oC; Fritz

1965; Schaefer 1967; Horn 1970a) and euryhaline (5-32

ppt; Musick 1972).

EGGS

Butterfish eggs are buoyant, transparent, and

spherical (0.68-0.82 mm diameter; Wheatland 1956;

Colton and Marak 1969; Martin and Drewry 1978; Elliott

and Jiminez 1981). The incubation period is about 48 hrs

at 18oC; 50% of eggs hatched at 72 hrs at about 15oC

(Martin and Drewry 1978; Colton and Honey 1963).

Eggs have been collected between 12.8-22.5oC and 78-

100% seawater (Martin and Drewry 1978). At hatching,

butterfish are 1.68-1.75 mm; yolk absorption is complete

by 2.48-2.64 mm (Colton and Honey 1963; Colton and

Marak 1969).

LARVAE

Butterfish larvae range from 2.6 to 16 mm standard

length (SL) (Martin and Drewry 1978). By 6 mm they

have the thin, deep body that is characteristic of adults

and by 15-16 mm they have a forked tail (Horn 1970a;

Ditty and Truesdale 1983). At 10-15 mm, larvae are more

nektonic than planktonic (Martin and Drewry 1978) and

are caught in neuston nets (Powles and Stender 1976; Lux

and Wheeler 1992). They begin to associate with

jellyfish, Sargassum, and other flotsam at this size

(Mansueti 1963; Haedrich 1967; Horn 1970b; Thomas

and Milstein 1973; Lippson and Lippson 1984). Larvae

may undertake diel vertical migrations; more butterfish

larvae were collected between 0-4 m at night than during

the day (Kendall and Naplin 1981). Metamorphosis is

gradual as the larvae progressively assume juvenile

characters (Able and Fahay 1998). Rotunno (1992)

reported growth rates of 0.227 mm/day for fish 6.0-28.0

mm SL based on otolith analyses.

JUVENILES

Juvenile butterfish range from 16 mm to about 120

mm SL (Martin and Drewry 1978). During their first

year, they grow to 76-127 mm, or about half their adult

size (Hildebrand and Schroeder 1928; Klein-MacPhee, in

review). Early-spawned individuals are 76-102 mm in the

fall; late-spawned individuals are 51-76 mm in the fall

and 76-127 mm the following spring (Martin and Drewry

1978). Young butterfish (< 30 mm) often live in the

shelter of large jellyfishes during their first summer.

Although this commensal association is not essential, it is

a source of food and provides young butterfish some

protection from their predators (Mansueti 1963; Horn

1970b, 1975).

ADULTS

Adult butterfish range from about 120 mm to 305

mm SL (Hildebrand and Schroeder 1928) with an average

length of 150-230 mm (Klein-MacPhee, in review). The

median length at maturity (L50) for butterfish collected on

the northeast shelf (1986-1989) was 12.0 cm total length

(TL) for females and 11.4 cm TL for males (O’Brien et

al. 1993), which corresponds to an age of about 1 year

(Horn 1970a; DuPaul and McEachran 1973). In

Chesapeake Bay, butterfish begin to mature during their

second summer (age 1) and most individuals are mature

by their third summer (DuPaul and McEachran 1973). In

the New York Bight, ripe females 124-242 mm FL were

collected in 3-145 m of water from May through August;

less than 5% of the ripe females were collected in the

Hudson-Raritan estuary (Wilk et al. 1990). At 2+ years

of age, butterfish are about 17 cm and at 3+, they are

about 19 cm (Waring and Murawski 1982).

REPRODUCTION

Butterfish are broadcast spawners (Horn 1970a) and

spawn primarily in the evening or at night (Ferraro 1980;
Page 2

Kendall and Naplin 1981), but no direct observations

have been made (Klein-MacPhee, in review). Butterfish

may spawn in the upper part of the water column during

the evening; more eggs were collected between 0-4 m at

night in the Middle Atlantic Bight than during the day

(Kendall and Naplin 1981).

Butterfish are usually reported to spawn offshore

(e.g., Wang and Kernehan 1979). Butterfish may spawn a

few miles out to sea off Woods Hole, MA and return

inshore when they are spent (Klein-MacPhee, in review).

However, eggs and larvae have been collected in coastal

waters and most estuaries in the northern part of the

Middle Atlantic Bight (Hildebrand and Schroeder 1928;

Herman 1963; Martin and Drewry 1978; Lux and

Wheeler 1992; Able and Fahay 1998). Early stage eggs

have been collected in Narragansett Bay and Salem

Harbor (Herman 1963; Bourne and Govoni 1988; Elliott

and Jiminez 1981), Raritan Bay, NJ (Croker 1965), and in

the lower portions of Chesapeake Bay (Lippson and

Moran 1974), but not in Delaware Bay (Wang and

Kernehan 1979).

Water temperatures appear to regulate butterfish

reproduction as spawning dates are progressively later in

the year in the northern part of its range (Murawski et al.

1978; Rotunno and Cowen 1997; Able and Fahay 1998).

Spawning may occur year round in the South Atlantic

Bight with a peak in spring (Fahay 1975; Able and Fahay

1998). Spawning probably does not occur below 15oC

(Colton 1972).

Butterfish begin spawning in Chesapeake Bay as

early as late May with a peak in activity in June and July

(Hildebrand and Schroeder 1928; Pearson 1941).

Spawning in the Middle Atlantic Bight occurs from May

through October (Smith et al. 1980); the gonad weight of

fish > 15 cm increases in March and April, reaches its

maximum during June and July, and decreases in the fall

(Kawahara 1977b). In Long Island Sound, spawning

occurs from June through late August with a peak in late

July; the principal spawning areas are in the eastern part

of the sound (Perlmutter 1939). In Narragansett Bay,

butterfish eggs are found from June to August (Herman

1963). In Massachusetts Bay, butterfish spawn from June

to August (Bigelow and Schroeder 1953). In the Gulf of

Maine, spawning begins in May-June, peaks in July, and

ends in August (Bigelow and Schroeder 1953; Smith et al.

1980). On the Scotian Shelf, spawning occurs from July

to October (Markle and Frost 1985).

The spawning period may be more protracted in the

Middle Atlantic Bight than previously thought. Rotunno

(1992) and Rotunno and Cowen (1997) estimated

spawning times from a birthdate analysis of otoliths from

butterfish up to about 50 mm SL collected in the Middle

Atlantic and South Atlantic bights. Spawning began in

February and continued through at least late July. It

began in the south and progressed northward over time,

which is consistent with the temporal and spatial

distribution of larvae, and suggests that butterfish spawn

as they migrate north and inshore on their annual

migration in association with seasonal warming of waters

on the northeast shelf.

FOOD HABITS

Butterfish feed mainly on planktonic prey including

thaliaceans (primarily Larvacea and Hemimyaria),

mollusks (primarily squids), crustaceans (copepods,

amphipods, and decapods), coelenterates (primarily

hydrozoans), polychaetes (primarily Tomopteridae and

Goniadidae), small fishes, and ctenophores (Fritz 1965;

Leim and Scott 1966; Haedrich 1967; Horn 1970a, b;

Schreiber 1973; Mauer and Bowman 1975; Oviatt and

Kremer 1977; Tibbets 1977; Murawski et al. 1978;

Bowman and Michaels 1984; Klein-MacPhee, in review).

The food habits of butterfish collected during the

northeast shelf during Northeast Fisheries Science Center

(NEFSC) bottom trawl surveys [see Reid et al. (1999) for

details] were similar to diets reported in the literature

(Figure 3). The stomach contents were dominated by

unidentifiable animal remains. Arthropods dominated the

identifiable items, followed by urochordates (thaliaceans

and larvaceans), unidentified plankton, annelids (probably

polychaetes), chaetognaths (arrowworms), mollusks

(probably squids), cnidarians (coelenterates, probably

jellyfish), and fishes.

PREDATION

Butterfish are preyed on by many species including

haddock, silver hake, goosefish, weakfish, bluefish,

swordfish, sharks (hammerhead), and longfin inshore

squid (Bigelow and Schroeder 1953; Scott and Tibbo

1968; Horn 1970a; Maurer and Bowman 1975; Tibbets

1977; Stillwell and Kohler 1985; Brodziak 1995; KleinMacPhee, in review).

MIGRATION

North of Cape Hatteras, butterfish have a seasonal

inshore-offshore north-south migration in response to

changing water temperatures. There is a limited seasonal

inshore-offshore migration south of Cape Hatteras

(Caldwell 1961; Fritz 1965; Horn 1970a; Klein-MacPhee,

in review). During the summer, butterfish move north

and inshore to feed on planktonic fish, squid, crustaceans,

and jellyfish, and to reproduce. They remain near the

surface at depths of 22-55 m and often come close

inshore; schools are frequently seen on shallow flats and

in sheltered bays and estuaries (Bigelow and Schroeder

1953; Klein-MacPhee, in review).

Butterfish are common in the lower Chesapeake Bay

from March through November (Geer and Austin 1997;
Page 3

Murdy et al. 1997). They occur in Great Bay, NJ and

nearby coastal waters from June through November (Able

and Fahay 1998) and in the surf zone off Long Island

from June through October (Schaefer 1967). They appear

off Rhode Island by the last half of April and off Woods

Hole, MA by mid-May, although they are not abundant

there until June. Butterfish appear on Georges Bank in

early June, but are not abundant until late June or early

July. They occur in the Gulf of Maine from late Juneearly July through the fall (Bigelow and Schroeder 1953;

Overholtz and Tyler 1985; Klein-MacPhee, in review).

They are found in New Hampshire waters from July to

October with a peak in abundance in September

(MAFMC 1995). Butterfish are common along the coast

of Maine and, in some years, they are common along the

coast of Nova Scotia bordering the Gulf of Maine

(Bigelow and Schroeder 1953).

During the winter, the stock moves south and

offshore. Butterfish are found near the bottom over sand,

mud, and rock bottoms. They have been caught to about

200 m deep in the northwest Atlantic (Bigelow and

Schroeder 1953; Klein-MacPhee, in review) and over 350

m in the South Atlantic Bight (Barans and Burrell 1976).

Butterfish are absent from nearshore waters off New

Jersey from January through late April (Milstein 1974;

Milstein and Hamer 1976). South of Delaware Bay, the

winter offshore movement is not so extensive and some

individuals move south in shallow water (Waring and

Murawski 1982).

STOCK STRUCTURE

Butterfish range from Newfoundland to Florida and

are considered a unit stock (Brodziak 1995; KleinMacPhee, in review). There may be two stocks south of

Cape Hatteras that are isolated by depth, although the

shallow stock (< 20 m) may be a Peprilus triacanthusPeprilus burti hybrid (Caldwell 1961; Horn 1970a; KleinMacPhee, in review) or P. burti, a Gulf of Mexico species

(Pershbacher et al. 1979).

HABITAT CHARACTERISTICS

Butterfish are pelagic fishes that form loose schools,

often near the surface (Schreiber 1973; Dery 1988;

Brodziak 1995). They winter near the edge of the

continental shelf in the Middle Atlantic Bight and migrate

inshore in the spring into southern New England and Gulf

of Maine waters. During the summer, butterfish occur

over the entire Mid-Atlantic shelf from sheltered bays and

estuaries out to about 200 m. In late fall, butterfish move

southward and offshore in response to falling water

temperatures (Fritz 1965; Horn 1970a; Schreiber 1973;

Waring 1975; Azarovitz et al. 1980; Klein-MacPhee, in

review).

Table 1 summarizes the environmental conditions

where butterfish eggs, larvae, juveniles, and adults have

been collected based on a literature survey and analyses

of several fishery-independent databases [see Reid et al.

(1999) for survey methods and location maps].

EGGS AND LARVAE

Butterfish eggs and larvae are pelagic and occur from

the outer continental shelf to the lower, high salinity parts

of estuaries in Middle Atlantic Bight. Eggs have been

collected between 12-23oC and larvae have been collected

between 4-28oC; eggs and larvae occur at salinities that

range from estuarine to full strength seawater (Table 1).

Larvae may undertake diel vertical migrations (Kendall

and Naplin 1981). Larger larvae (10-15 mm) are more

nektonic than planktonic; larger larvae and pelagic

juveniles (< 30 mm) often associate with jellyfish,

Sargassum, and other flotsam (Mansueti 1963; Haedrich

1967; Horn 1970b; Thomas and Milstein 1973; Lippson

and Lippson 1984).

Eggs were collected during the NEFSC Marine

Resources Monitoring, Assessment and Prediction

program (MARMAP) ichthyoplankton survey at water

temperatures ranging from 6o to 26oC; most eggs were

collected between 11-17oC (Figure 4). Eggs were

collected in surface waters (upper 200 m or within 5 m of

bottom where station depths were < 200 m) in depths

ranging from 10 to 1250 m (Figure 4). Most eggs were

collected in water depths < 200 m.

Larvae were collected during the MARMAP

ichthyoplankton survey at water temperatures ranging

from 7-26oC; most larvae were collected at 9-19oC

(Figure 5). Larvae were collected in surface waters in

depths ranging from 10 to 1750 m; most larvae were

collected in water depths < 120 m (Figure 5).

Eggs and larvae are common in the high salinity

zones of some estuaries in southern New England and the

Middle Atlantic Bight and in the mixing zone in

Chesapeake Bay (Table 2a).

JUVENILES AND ADULTS

Juvenile and adult butterfish are pelagic fishes that

form loose schools, often near the surface (Schreiber

1973; Dery 1988; Brodziak 1995). They are eurythermal

(4.4-21.6oC) and euryhaline (5-32 ppt) and are frequently

found over sand, mud, and mixed substrates (Table 1). In

Long Island Sound, butterfish were collected less

frequently at low dissolved oxygen levels (2.0-2.9ml/l)

(Howell and Simpson 1994).

During the summer, butterfish occur inshore where

they remain near the surface; schools are frequently seen

on shallow flats and in sheltered bays, estuaries, and the

surf zone (Bigelow and Schroeder 1953; Leim and Scott
Page 4

1966; Schaefer 1967; Klein-MacPhee, in review).

Smaller juveniles often aggregate under floating objects

including the bells of coelenterates (Pearson 1941;

Bigelow and Schroeder 1953; Mansueti 1963; Haedrich

1967; Horn 1970b, 1975; Lippson and Moran 1974;

Milstein 1974; Scott and Scott 1988). Larger juveniles

are pelagic schooling fishes that may congregate near the

bottom during the day and disperse upwards at night

(Waring 1975).

Juvenile and adult butterfish are common to abundant

in the high salinity and mixing zones of estuaries from

Massachusetts Bay to the mid-Atlantic; they are rare to

uncommon in the high salinity and mixing zones of

estuaries in the central and northern Gulf of Maine and in

the South Atlantic Bight (Tables 2a, b). In the Gulf of

Maine and Middle Atlantic Bight, butterfish move

offshore during the winter; fish are found near the bottom

over sand, mud, and rock substrates (Bigelow and

Schroeder 1953; Klein-MacPhee, in review). The

offshore migration is not as pronounced south of

Delaware Bay where winter water temperatures are

warmer (Waring and Murawski 1982). In the South

Atlantic Bight, butterfish are present throughout most of

the year in nearshore waters (Keiser 1976).

In the NEFSC bottom trawl survey (1963-1997),

juvenile and adult butterfish were collected on the

continental shelf from 10 m of water nearshore out to

about 360 m of water offshore; most juveniles and adults

were collected in water depths < 180 m (Figure 6).

Adults were distributed somewhat deeper than juveniles

in all seasons. Bottom-water temperatures where

juveniles and adults were captured ranged from 3o to

28oC; most fish were collected between 7-20oC (Figure

6). Modal water temperatures during spring and fall

surveys were 10-14oC for juveniles and adults.

In the Massachusetts trawl survey (1978-1996),

juvenile and adult butterfish were collected at depths

ranging from 5 to 80 m; most juveniles were collected

between 10-35 m and most adults between 10-50 m

(Figure 7). Bottom water temperatures ranged from 9-

15oC in the spring and 7-22oC in the fall (Figure 7).

Adults were caught deeper than juveniles in the fall when

water temperatures were lower.

In the Rhode Island Narragansett Bay/Coastal trawl

survey, juvenile and adult butterfish were collected at

depths between 10-120 ft (3-37 m); most juveniles and

adults were collected between 30-110 ft (10-34 m).

Bottom water temperatures for juveniles and adults at the

time of collection ranged from 9-24oC in the summer and

fall and 5-15oC in the winter and spring.

In the Connecticut Long Island Sound trawl survey,

juvenile and adult butterfish were collected at depths

between 6-60 m; most fish were collected between 10-30

m. Bottom water temperatures for juveniles and adults at

the time of collection ranged from 7-18oC in the spring

and 8-23oC in the fall; most fish were captured at 9-15oC

in the spring and 16-21oC in the fall. Bottom water

salinities at the time of collection ranged from 18-32 ppt;

most fish were captured at 26-29 ppt.

In the Hudson-Raritan trawl survey, juvenile and

adult butterfish were collected at depths ranging from 10-

75 ft (3-23 m) (Figure 8). Bottom water temperatures

ranged from 8-26oC, salinities ranged from 19-32 ppt, and

dissolved oxygen ranged from 3-10 mg/l (Figure 8).

GEOGRAPHICAL DISTRIBUTION

Butterfish range from Newfoundland and the Gulf of

St. Lawrence to the Atlantic and Gulf coasts of Florida

(Figure 2), but they are most abundant from the Gulf of

Maine to Cape Hatteras (Haedrich 1967; Horn 1970a;

Powell et al. 1972; Cooley 1978; Scott and Scott 1988;

Brodziak 1995; Klein-MacPhee, in review). Butterfish

spend the winter near the edge of the continental shelf in

the Middle Atlantic Bight and migrate inshore in spring to

waters off southern New England and into the Gulf of

Maine. During the summer, butterfish range from the

Gulf of Maine to the South Atlantic Bight where they are

found from sheltered bays and estuaries (Table 3) across

the shelf to depths of 200 m and greater. In late fall,

butterfish move southward and offshore in response to

falling water temperatures (Fritz 1965; Horn 1970a;

Schreiber 1973; Waring 1975; Azarovitz et al. 1980;

Klein-MacPhee, in review). During the winter, they are

largely absent from bays and estuaries in the Middle

Atlantic Bight and Gulf of Maine (Table 3).

EGGS

Butterfish eggs have been reported in the Gulf of

Maine, on Georges Bank, in the Middle Atlantic Bight,

and off North Carolina (Smith et al. 1980; Rotunno 1992;

MAFMC 1995; Rotunno and Cowen 1997). They have

also been collected in Salem Harbor, MA and

Narragansett Bay, RI (Herman 1963; Bourne and Govoni

1988; Elliott and Jiminez 1981), Block Island Sound

(Merriman and Sclar 1952), Long Island Sound

(Wheatland 1956), Peconic Bay, NY (Ferraro 1980),

Raritan Bay, NJ (Croker 1965), and Chesapeake Bay

(Lippson and Moran 1974).

During the MARMAP ichthyoplankton survey,

butterfish eggs were collected from Cape Hatteras to the

northern Gulf of Maine from April through September

(Figure 9). Eggs first appeared in ichthyoplankton

collections in April; by May, eggs were distributed along

the edge of the continental shelf between Cape Hatteras

and Georges Bank and inshore in the southern and middle

Mid-Atlantic Bight. As water temperatures increased on

the shelf, eggs were found progressively closer to the

coast from south to north. Eggs were most abundant and

most frequently encountered in July; they were most

abundant in the Gulf of Maine in August. By September,
Page 5

egg abundance declined dramatically; no eggs were

collected from October to March.

In coastal bays and estuaries, butterfish eggs were

recorded as far north as Penobscot Bay and as far south as

Chesapeake Bay (Stone et al. 1994). Eggs were abundant

in Narragansett Bay and common in Massachusetts Bay,

Cape Cod Bay, Waquoit Bay, Buzzards Bay, Long Island

Sound, Gardiners Bay, Great South Bay, and Chesapeake

Bay (Table 2a).

LARVAE

Butterfish larvae have been reported from the New

York Bight and Georges Bank (Smith et al. 1980; Wilk et

al. 1990; Rotunno 1992; MAFMC 1995; Rotunno and

Cowen 1997), in Buzzards Bay, MA (Lux and Wheeler

1992), Narragansett Bay, RI (Herman 1963; Bourne and

Govoni 1988; Elliott and Jiminez 1981), Raritan Bay, NJ

(Croker 1965), Great Bay, NJ (Able and Fahay 1998),

Chesapeake Bay (Lippson and Moran 1974), and in the

South Atlantic Bight as far south as Cape Kennedy, FL

(Fahay 1975; Powles and Stender 1976; Rotunno 1992;

Rotunno and Cowen 1997). Larvae were not abundant in

the South Atlantic Bight (< 0.5% of total

ichthyoplankton) and did not occur frequently (< 10% of

stations in a survey of 73 coastal stations) (Fahay 1975).

During the MARMAP ichthyoplankton survey,

butterfish larvae were collected from Cape Hatteras into

the Gulf of Maine in every month except December

(Figure 10). Larvae first appeared in ichthyoplankton

collections in January. From January through April,

larvae were collected primarily off Cape Hatteras. In

May and June, larvae began to appear along the edge of

the continental shelf between Cape Hatteras and Georges

Bank and inshore in the southern portion of the Middle

Atlantic Bight. As water temperatures increased on the

shelf, larvae were found progressively closer to the coast

from south to north. Larvae were most abundant and

most frequently encountered in July and August across

the continental shelf in the Middle Atlantic Bight

northward to Georges Bank. The abundance of larvae

declined sharply from September through November.

In the coastal bays and estuaries of New England and

the mid-Atlantic, butterfish larvae were recorded as far

north as Penobscot Bay and as far south as Chesapeake

Bay (Stone et al. 1994). Larvae were common in Boston

Harbor, Waquoit Bay, Buzzards Bay, Narragansett Bay,

Long Island Sound, Gardiners Bay, Great South Bay,

Great South Bay, and Chesapeake Bay (Table 2a).

JUVENILES

Juvenile butterfish occur from Nova Scotia to the

Atlantic and Gulf coasts of Florida, but they are most

abundant from the Gulf of Maine to Cape Hatteras

(Bigelow and Schroeder 1953; Haedrich 1967; Horn

1970a; Powell et al. 1972; Cooley 1978; Scott and Scott

1988; Brodziak 1995; Klein-MacPhee, in review). They

occur in the high salinity and mixed salinity zones of most

estuaries from the Gulf of Maine to Florida (Table 2a)

(Jury et al. 1994; Stone et al. 1994; Geer and Austin

1997; Murdy et al. 1997).

During the NEFSC Bottom trawl survey, juvenile

butterfish were collected from the northern Gulf of Maine

south to Cape Lookout, South Carolina (Figure 11).

During the winter and spring, juveniles were collected

along the outer continental shelf from southern New

England to Cape Hatteras and along the coast near Cape

Hatteras. During the summer, juvenile butterfish were

collected near the coast throughout the Middle Atlantic

Bight and on Georges Bank. During the fall, they were

abundant across the shelf throughout the Middle Atlantic

Bight and on Georges Bank.

Juvenile butterfish were collected in spring and fall

by the Massachusetts Trawl Survey, but catches were 1-2

orders of magnitude greater in the fall (Figure 12).

During the Southeast Area Monitoring and Assessment

Program-South Atlantic (SEAMAP-SA) bottom trawl

survey, juvenile butterfish were collected from Cape

Lookout, South Carolina to Cape Kennedy, Florida

(Figure 13). Catches were smallest during the winter and

largest during the summer.

In the coastal bays and estuaries of New England and

the mid-Atlantic, juvenile butterfish were recorded from

Passamaquoddy Bay, Maine south to the James River in

Virginia (Table 2a) (Stone et al. 1994). South of Cape

Hatteras, juveniles occurred in bays and estuaries in South

Carolina, Georgia, and Florida (Table 2a). Juveniles were

abundant in Buzzards Bay, Narragansett Bay, and Long

Island Sound, and common in most of the remaining bays

and estuaries between Massachusetts Bay and Chesapeake

Bay.

In Narragansett Bay, juvenile butterfish were

collected in all seasons, but they were rare in winter and

spring; they were most abundant in summer when they

occurred throughout the bay (Figure 14). In Long Island

Sound, butterfish appeared in May; abundance peaked in

September-October and declined in November (Figure

15). Juveniles composed 17% of all butterfish caught in

May, 91% in September-October, and 73% in November.

Juveniles appear in surf zone off Long Island in July and

are common from August through October (Schaefer

1967). In the Hudson-Raritan estuary, juveniles were

caught in trawls from spring through fall (Figure 16).

ADULTS

Adult butterfish have been reported from

Newfoundland to the Atlantic and Gulf coasts of Florida,

but they are most abundant from the Gulf of Maine to

Cape Hatteras (Bigelow and Schroeder 1953; Haedrich
Page 6

1967; Horn 1970a; Powell et al. 1972; Cooley 1978; Scott

and Scott 1988; Brodziak 1995; Klein-MacPhee, in

review). They have been collected in high salinity and

mixed salinity zones of most estuaries from the Gulf of

Maine to Florida (Tables 2a, b) (Hildebrand and

Schroeder 1928; DuPaul and McEachran 1973; Wilk and

Silverman 1976b; Jury et al. 1994; Stone et al. 1994; Geer

and Austin 1997; Murdy et al. 1997).

During the NEFSC bottom trawl survey, adult

butterfish were collected from the northern Gulf of Maine

south to below Cape Lookout, South Carolina (Figure 11).

During the winter and spring, they were distributed along

the outer continental shelf from southern New England to

Cape Hatteras; they occurred along the coast from Cape

Hatteras to Maryland. During the summer, adult

butterfish were collected across the shelf throughout the

Middle Atlantic Bight, on Georges Bank, and in the

coastal Gulf of Maine. During the fall, they were

abundant on the shelf throughout the Middle Atlantic

Bight, on Georges Bank, and in Massachusetts Bay.

In the Massachusetts Trawl Survey, adult butterfish

were collected in the spring primarily south of Cape Cod

and in Buzzards Bay, and in the fall primarily in Buzzards

Bay, Massachusetts Bay, and around Cape Ann (Figure

12). During the SEAMAP-SA bottom trawl survey, adult

butterfish were collected from Cape Lookout, South

Carolina to Cape Kennedy, Florida (Figure 13). The size

of the catches was similar throughout the year. Butterfish

are present in nearshore waters off South Carolina

throughout most of the year (Keiser 1976).

In the coastal bays and estuaries of New England and

the mid-Atlantic, adult butterfish were recorded from

Passamaquoddy Bay in Maine south to the James River in

Virginia (Jury et al. 1994; Stone et al. 1994). South of

Cape Hatteras, adults occurred in bays and estuaries in

South Carolina, Georgia, and Florida (Table 2a, b).

Adults were abundant in Buzzards Bay, Narragansett Bay,

and Long Island Sound, and common in most of the

remaining bays and estuaries between Massachusetts Bay

and Chesapeake Bay (Table 2b). Spawning adults were

recorded from Massachusetts Bay south to the

Chesapeake Bay, but were common only in Long Island

Sound, Gardiners Bay, Great South Bay, and Chesapeake

Bay (Table 2b).

In Narragansett Bay, adult butterfish were collected

in all seasons, but they were rare in winter and spring;

they were most abundant in summer when they occurred

throughout the bay (Figure 14). In Long Island Sound,

butterfish appeared in May; abundance peaked in

September-October and declined in November (Figure 15;

Wheatland 1956). Adults composed 83% of all butterfish

caught in May, 9% in September-October, and 27% in

November. Adults appear in the surf zone off Long

Island in May and are common from June through

October (Schaefer 1967). Butterfish were among the

most abundant species in both of these Long Island

surveys. In the Hudson-Raritan estuary, adults were

caught from spring through fall (Figure 16).

STATUS OF THE STOCKS

A fishery for butterfish has existed since the late

1800s (Murawski and Waring 1979); from 1920 to 1962,

the average annual landings in US waters were 3,000 mt

(Waring 1975). In 1963, distant water fleets from Japan,

Poland, and the USSR began targeting butterfish from late

autumn through early spring when the fish were

concentrated offshore (Murawski and Waring 1979;

MAFMC 1995). Annual landings increased to a record

19,500 mt in 1973 (Figure 17) (Brodziak 1995).

Restrictions were placed on the foreign fisheries and

landings subsequently decreased to an average of 6,100

mt from 1977 to 1987. Directed foreign fishing was

halted in 1987 and landings continued to decline to an

average 2,500 mt in the domestic fishery from 1987 to

1992 (Brodziak 1995; MAFMC 1995). The domestic

fishery targeted butterfish from late spring through fall in

inshore areas (Murawski and Waring 1979). Butterfish

landings totaled 4,500 mt in 1993 and came primarily

from southern New England (79% in Rhode Island ports)

and the New York Bight. These landings were 60%

higher than landings in 1992 and were comparable with

record domestic catches in 1987 (Brodziak 1995).

Butterfish biomass estimated from the Northeast

Fisheries Science Center bottom trawl surveys has made

several record lows and near record highs in the last

decade (Figure 17). Despite seasonal increases in

biomass and pre-recruit indices, butterfish stock size has

decreased and commercial landings remain low

(Northeast Fisheries Science Center 1994). Although the

demand for butterfish has declined in recent years, the

capacity for increased landings remains in an underexploited fishery (Brodziak 1995). The butterfish stock is

not overfished nor approaching an overfished condition

(National Marine Fisheries Service 1997).

ACKNOWLEDGMENTS

The authors thank C. Steimle, J. Berrien, and R.

Ramsey-Cross for help with the literature review and B.

Collette for providing the draft section on butterfish from

the update of Bigelow and Schroeder (1953). W. Morse

produced maps and histograms for the Massachusetts,

Rhode Island, Connecticut, and SEAMAP-SA surveys. S.

Wilk produced the maps and histograms for the HudsonRaritan survey. J. Vitaliano produced the gut contents

figure from the NEFSC food habits database. L.

Cargnelli, S. Griesbach, and C. Noonan assisted with

editing and formatting.
Page 7

REFERENCES CITED

Able, K.W., and M.P. Fahay. 1998. The first year in the

life of estuarine fishes in the Middle Atlantic Bight.

Rutgers University Press, New Brunswick, NJ. 342 p.

Austin, H.M. 1973. The ecology of Lake Montauk:

planktonic fish eggs and larvae. New York Ocean

Sci. Lab. Tech. Rep. No. 21. Montauk, NY.

Austin, H.M. 1976. Distribution and abundance of

ichthyoplankton in the New York Bight during the

fall in 1971. N.Y. Fish Game J. 23: 58-72.

Azarovitz, T.R., C.J. Byrne, E.S. Bevacqua, L.I. Despres,

and H.A. Foster. 1980. Distribution and abundance

trends of 22 selected species in the Middle Atlantic

Bight from bottom trawl surveys during 1967-1979:

Final report to the U.S. Minerals Management

Service. National Marine Fisheries Service, Northeast

Fisheries Center, Woods Hole, MA. 581 p.

Barans, C.A. and V.G. Burrell, Jr. 1976. Preliminary

findings of trawling on the continental shelf off the

southeastern United States during four seasons

(1973-1975). Tech. Rep. No. 13. South Carolina

Marine Resources Center, South Carolina Wildlife

and Marine Resources Dep., Charleston, SC. 16 p.

Berrien, P.L., M.P. Fahay, A.W. Kendall Jr., and W.G.

Smith. 1978. Ichthyoplankton from the RV Dolphin

survey of continental shelf waters between Martha’s

Vineyard, Massachusetts and Cape Lookout, North

Carolina, 1965-1966. U.S. Natl. Mar. Fish. Serv.,

Northeast Fish. Cent., Sandy Hook Lab. Tech. Ser.

Rep. 15. 152 p.

Bigelow, H. B. and W.C. Schroeder. 1953. Fishes of the

Gulf of Maine. U.S. Fish Wildl. Serv. Fish. Bull. 53.

577 p.

Bourne, D.W. and J.J. Govoni. 1988. Distribution of fish

eggs and larvae and patterns of water circulation in

Narragansett Bay, 1972-1973. In M.P. Weinstein ed.

Larval fish and shellfish transport through inlets. p.

132-148. Am. Fish. Soc. Symp. 3. American

Fisheries Society, Bethesda, MD.

Bowman, R.E. and W.L. Michaels. 1984. Food of

seventeen species of northwest Atlantic fish. NOAA

Tech. Mem. NMFS-F/NEC-28. 183 p.

Brodziak, J. 1995. Atlantic butterfish. In Conservation

and Utilization Division, Northeast Fisheries Science

Center eds. Status of the fishery resources off the

northeastern United States for 1994. p. 102-103.

NOAA Tech. Mem. NMFS-F/NEC-108.

Caldwell, D.K. 1961. Populations of butterfish, Peprilus

triacanthus, with systematic comments. Bull. S.

Calif. Acad. Sci. 60: 19-31.

Colton, J.B., Jr. 1972. Temperature trends and the distribution of groundfish in continental shelf waters,

Nova Scotia to Long Island. Fish. Bull. (U.S.) 70:

637-658.

Colton, J.B., Jr. and K.A. Honey. 1963. The eggs and

larval stages of the butterfish, Poronotus triacanthus.

Copeia 2: 447-450.

Colton, J.B., Jr., and R.R. Marak. 1969. Guide for

identifying the common planktonic fish eggs and

larvae of continental shelf waters, Cape Sable to

Block Island. U.S. Bur. Comm. Fish., Woods Hole,

Lab. Ref. Doc. No. 69-9. 43 p.

Colton, J.B., Jr., W.G. Smith, A.W. Kendall Jr., P.L.

Berrien, and M.P. Fahay. 1979. Principal spawning

areas and times of marine fishes from Cape Sable to

Cape Hatteras. Fish. Bull. (U.S.) 76: 911-915.

Cooley, N.R. 1978. An inventory of the estuarine fauna in

the vicinity of Pensacola, Florida. Florida Mar. Res.

Pub. No. 31. Florida Dep. Nat. Resources, Mar. Res.

Lab. St. Petersburg, FL.119 p.

Croker, R.A. 1965. Planktonic fish eggs and larvae of

Sandy Hook estuary. Chesapeake Sci., 6: 92-95.

Dahlberg, M.D. 1972. An ecological study of Georgia

coastal fishes. Fish. Bull. (U.S.) 70: 323-353.

Dery, L.M. 1988. Butterfish, Peprilus triacanthus. In J.

Penttila and L.M. Dery eds. Age determination

methods for northwest Atlantic species. p. 85-92.

NOAA Tech. Rep. NMFS-72.

Ditty, J.G. and F.M. Truesdale. 1983. Comparative larval

development of Peprilus burti, P. triacanthus and P.

paru (Pisces: Stromateidae) from the western North

Atlantic. Copeia 2: 397-406.

Draganik, B. and C. Zukowski. 1966. The rate of growth

of butterfish (Poronotus triacanthus (Peck)) and

ocean pout (Macrozoarces americanus (Bloch and

Schneider)) from the region of Georges Bank. Int.

Comm. Northwest Atl. Fish. (ICNAF) Res. Doc. 66-

42.

DuPaul, W.D. and J.D. McEachran. 1973. Age and

growth of butterfish, Peprilus triacanthus in the

lower York River. Chesapeake Sci. 14: 205-207.

Edwards, R.L., R. Livingstone Jr., and P.E. Hamer. 1962.

Winter water temperatures and an annotated list of

fishes - Nantucket Shoals to Cape Hatteras. Albatross

III Cruise No. 126. U.S. Fish Wildl. Serv. Spec. Sci.

Rep. Fish. 397. 31 p.

Elliott, E.M. and D. Jimenez. 1981. Laboratory manual

for the identification of ichthyoplankton from the

Beverly-Salem Harbor area. Massachusetts Dep.

Fish., Div. Mar. Fish. Boston, MA. 230 p.

Fahay, M.P. 1975. An annotated list of larval and juvenile

fishes captured with surface-towed meter net in the

South Atlantic Bight during four RV Dolphin cruises

between May 1967 and February 1968. NOAA Tech.

Rep. NMFS-SSRF-685. 39 p.

Ferraro, S.P. 1980. Daily time of spawning of 12 fishes in

the Peconic Bays, New York. Fish. Bull. (U.S.) 78:

455-464.

Fritz, R.L. 1965. Autumn distribution of groundfish

species in the Gulf of Maine and adjacent waters,

1955-1961. Serial Atlas of the Marine Environment.

Folio 10. American Geographical Society, NY. 48 p.
Page 8

Geer, P.J. and H.M Austin. 1997 Estimation of relative

abundance of recreationally important finfish in the

Virginia portion of Chesapeake Bay. Ann. Prog.

Rep., Virginia Inst. Mar. Sci., Gloucester Point, VA.

153 p. + appendices.

Goode, G.B. 1884. The fisheries and fishery industries of

the United States. Section I: Natural history of useful

aquatic animals. Govt. Print. Office, Washington,

DC. Plates.

Haedrich, R.L. 1967. The stromateoid fishes: systematics

and a classification. Bull. Mus. Comp. Zool. 135: 31-

139.

Herman, S.S. 1963. Planktonic fish eggs and larvae of

Narragansett Bay. Limnol. Oceanogr. 8: 103-109.

Hildebrand, S.F., and W.C. Schroeder. 1928. Fishes of

Chesapeake Bay. Bull. U.S. Bur. Fish. 43(1): 366 p.

Horn, M.H. 1970a. Systematics and biology of the

stromateiod fishes of the genus Peprilus. Bull. Mus.

Comp. Zool. Harv. Univ. 140: 165-262.

Horn, M.H. 1970b. The swim bladder as a juvenile organ

in stromateoid fishes. Breviora 359: 1-9.

Horn, M.H. 1975. Swim-bladder state and structure in

relation to behavior and mode of life in stromateoid

fishes. Fish. Bull. (U.S.) 73: 95-109.

Howell, P. and D. Simpson. 1994. Abundance of marine

resources in relation to dissolved oxygen in Long

Island Sound. Estuaries 17: 394-402.

Jury, S.H., J.D. Field, S.L. Stone, D.M. Nelson, and M.E.

Monaco. 1994. Distribution and abundance of fishes

and invertebrates in North Atlantic estuaries. ELMR

Rep. No. 13. NOAA/NOS Strategic Environmental

Assessments Division, Silver Spring, MD. 221 p.

Kawahara, S. 1977a. Age and growth of butterfish,

Poronotus triacanthus (Peck), in ICNAF Subarea 5

and Statistical Area 6. Int. Comm. Northwest Atl.

Fish. (ICNAF), Sel. Pap. 3: 73-78.

Kawahara, S. 1977b. Age and growth of butterfish,

Poronotus triacanthus (Peck), in ICNAF Subarea 5

and Statistical Area 6. Int. Comm. Northwest Atl.

Fish. (ICNAF) Res. Doc. 77/VI/27. 13 p.

Keiser, R.K., Jr. 1976. Species composition, magnitude

and utilization of the incidental catch of the South

Carolina shrimp fishery. Tech. Rep. No. 16, South

Carolina Marine Resources Center, South Carolina

Wildlife and Marine Resources Dep., Charleston, SC.

55 p. + appendices.

Kendall, A.W., Jr. and N.A. Naplin. 1981. Diel-depth

distribution of summer ichthyoplankton in the Middle

Atlantic Bight. Fish. Bull. (U.S.) 79: 705-726.

Klein-MacPhee, G. In review. Suborder Stromateoidei. In

B.B. Collette and G. Klein-MacPhee eds. Bigelow

and Schroeder’s fishes of the Gulf of Maine.

Smithsonian Institution Press, Washington, D.C.

Lang, P.J. 1974. Peprilus triacanthus (Peck), butterfish.

In Preoperational Ecological Monitoring Program of

Marine Environs of the Long Island Lighting Co.

(LILCO) Nuclear Power Generating Facility,

Shoreham, Long Island, NY. p. 169-176. Vol. 4.

Leim, A.H. and W.B. Scott. 1966. Fishes of the Atlantic

coast of Canada. Bull. Fish. Res. Board Can. 155.

485 p.

Lessard, J.A. 1974. Peprilus triacanthus (Peck), butterfish. Preoperational Ecological Monitoring Program

of Marine Environs of the Long Island Lighting Co.

(LILCO) Nuclear Power Generating Facility, Shoreham, Long Island, NY. p. 63-65 Vol. 2 (5) Ichthyoplankton.

Lippson, A.J. and R.L. Lippson. 1984. Life in the

Chesapeake Bay. Johns Hopkins University Press

Ltd., London, 219 p.

Lippson, A.J. and R.L. Moran. 1974. Manual for identification of early developmental stages of fishes of the

Potomac River Estuary. Environmental Technology

Center, Martin Marietta Corp., Baltimore, MD. 282

p.

Lux, F.E. and C.L. Wheeler. 1992. Larval and juvenile

fishes caught in a neuston survey of Buzzards Bay,

Massachusetts in 1979. U.S. Natl. Mar. Fish. Serv.,

Northeast Fish. Sci. Cent. Ref. Doc. 92-09. 12 p.

[MAFMC] Mid-Atlantic Fishery Management Council.

1995. Amendment #5 to the Fishery Management

Plan and the final environmental impact statement for

the Atlantic mackerel, squid, and butterfish fisheries.

August 1995. MAFMC. [Dover, DE.] 168 p. +

appendices.

Mansueti, R. 1963. Symbiotic behavior between small

fishes and jellyfishes, with new data on that between

the stromateoid, Peprilus alepidotus, and the

scyphomedusa, Chrysaora quinquecirrha. Copeia 1:

40-80.

Markle, D.F. and L.-A. Frost. 1985. Comparative morphology, seasonality, and a key to planktonic fish

eggs from the Nova Scotian shelf. Can. J. Zool. 63:

246-257.

Martin, F.D. and G.E. Drewry. 1978. Development of

fishes of the Mid-Atlantic Bight. An atlas of egg,

larval, and juvenile stages. Vol. 6: Stromateidae

through Ogcocephalidae. U.S. Fish Wildl. Serv. Biol.

Serv. Prog. FWS/OBS-78/12. 416 p.

Maurer, R.O., Jr. and R.E. Bowman. 1975. Food habits of

marine fishes of the northwest Atlantic - data report.

U.S. Natl. Mar. Fish. Serv., Northeast Fish. Cent.

Woods Hole Lab. Ref. Doc. No. 75-3. 90 p.

Merriman, D. and R.C. Sclar. 1952. The pelagic fish eggs

and larvae of Block Island Sound. In G.A. Riley et al.

eds. Hydrographic and biological studies of Block

Island Sound. p. 165-219. Bull. Bingham Oceanogr.

Collect. 13.

Milstein, C.B. 1974. Ocean trawl collections. In

Ecological studies in the bays and other waterways

near Little Egg Inlet and in the ocean in the vicinity

of the proposed site for the Atlantic Generating

Station, New Jersey. Progress report for the period

January through December 1973. p. 38-59. Prepared
Page 9

for Public Service Electric and Gas Co. by

Ichthyological Associates, Inc. Ithaca, NY.

Milstein, C.B. and D.P. Hamer. 1976. Fishes taken in the

vicinity of the site, the Great Bay-Mullica River

Estuary, and offshore with 25 ft trawl. In C.B.

Milstein ed. Ecological studies in the bays and other

waterways near Little Egg Inlet and in the ocean in

the vicinity of the proposed Atlantic Generating

Station, NJ. p. 21-42. Prepared for Public Service

Electric and Gas Co. by Ichthyological Associates,

Inc. Ithaca, NY.

Murawski, S.A., D.G. Frank, and S. Chang. 1978.

Biological and fisheries data on butterfish, Peprilus

triacanthus (Peck). U.S. Natl. Mar. Fish. Serv.,

Northeast Fish. Cent. Sandy Hook Lab. Tech. Ser.

Rep. No. 6. 38 p.

Murawski, S.A. and G.T. Waring. 1979. A population

assessment of butterfish, Peprilus triacanthus, in the

northwestern Atlantic Ocean. Trans. Am. Fish. Soc.

108: 427-439.

Murdy, E.O., R.S. Birdsong, and J.A. Musick. 1997.

Fishes of the Chesapeake Bay. Smithsonian Institution Press. Washington, DC. 324 p.

Musick, J.A. 1972. Fishes of Chesapeake Bay and

adjacent coastal plains. In M.L. Wass ed. A checklist

of the biota of the Lower Chesapeake Bay. p. 175-

212. Virginia Inst. Mar. Sci. Spec. Sci. Rep. 65. 290

p.

National Marine Fisheries Service. 1997. Report to

Congress. Status of fisheries of the United States:

Report on the status of fisheries of the United States.

September 1997. [Homepage of the National Marine

Fisheries Service]. [Online]. Available: http://www.

nmfs.gov/sfa/Fstatus.html.

Nelson, D.M., E.A. Irlandi, L.R. Settle, M.E. Monaco,

and L. Coston-Clements. 1991. Distribution and

abundance of fishes and invertebrates in southeast

estuaries. ELMR Rep. No. 9. NOAA/NOS Strategic

Environmental Assessments Division, Silver Spring,

MD. 167 p.

Northeast Fisheries Science Center. 1994. Report of the

17th Northeast Regional Stock Assessment Workshop. Stock Assessment Review Committee (SARC)

consensus summary of assessments. Northeast Fish.

Cent. Ref. Doc. 94-06. 124 p.

Obenchain, C.L. 1981. A study of the larval fish

community in the New York Bight, July 1974 to June

1976. In R. Lasker and K. Sherman. eds. The early

life history of fish: Recent studies. p. 217-219. Rapp.

P.-V. Reun. Cons. Int. Explor. Mer 178.

O’Brien, L., J. Burnett, and R.K. Mayo. 1993. Maturation

of nineteen species of finfish off the northeast coast

of the United States, 1985-1990. NOAA Tech Rep.

NMFS 113. 66 p.

Overholtz, W.J. and A.V. Tyler. 1985. Long-term responses of the demersal fish assemblages of Georges

Bank. Fish. Bull. (U.S.) 83: 507-520.

Oviatt, C.A. and P.M. Kremer. 1977. Predation on the

ctenophore, Mnemiopsis leidyi, by butterfish,

Peprilus triacanthus, in Narragansett Bay, Rhode

Island. Chesapeake Sci. 18: 236-240.

Pearson, J.C. 1941. The young of some marine fishes

taken in lower Chesapeake Bay, Virginia, with

special reference to the gray sea trout, Cynoscion

regalis (Bloch). Fish. Bull. (U.S.) 50: 79-102.

Perlmutter, A. 1939. A biological survey of the salt waters

of Long Island, 1938. An ecological survey of young

fish and eggs identified from the tow-net collections.

28th. Ann. Rep. New York State Cons. Dep., Suppl.,

Pt. II: 11-71.

Pershbacher, P.W., K.J. Sulak, and F.J. Schwartz. 1979.

Invasion of the Atlantic by Peprilus burti (Pisces:

Stromateidae) and possible implications Copeia

1979(3): 538-541.

Powell, D., L.M. Dwinell, and S.E. Dwinell. 1972. An

annotated listing of the fish reference collection at the

Florida Department of Natural Resources Marine

Research Laboratory. Spec. Sci. Rep. No. 36. Florida

Marine Research Laboratory, Department of Natural

Resources, St. Petersburg, FL. 179 p.

Powles, H. and B.W. Stender. 1976. Observations on

composition, seasonality and distribution of ichthyoplankton from MARMAP cruises in the South Atlantic Bight in 1973. Tech. Rep. No. 11. Marine Resources Research Inst., South Carolina Wildlife and

Marine Resources Dep., Charleston, SC. 47 p.

Reid, R., F. Almeida, and C. Zetlin. 1999. Essential fish

habitat source document: Fishery independent

surveys, data sources, and methods. NOAA Tech.

Mem. NMFS-NE-122. 39 p.

Rotunno, T.K. 1992. Species identification and temporal

spawning patterns of butterfish, Peprilus spp. in the

South and Mid-Atlantic Bights. M.S. thesis, State

University New York at Stony Brook, NY. 77 p.

Rotunno, T., and R.K. Cowen. 1997. Temporal and

spatial spawning patterns of the Atlantic butterfish,

Peprilus triacanthus, in the South and Middle

Atlantic Bights. Fish. Bull. (U.S.) 95: 785-799.

Schaefer, R.H. 1967. Species composition, size and

seasonal abundance of fish in the surf waters of Long

Island. N.Y. Fish Game J. 14: 1-46.

Schreiber, R.A. 1973. The fishes of Great South Bay.

M.S. thesis, State University of New York, Stony

Brook, NY. 199 p.

Schwartz, F.J., P. Perschbacher, M. McAdams, L. Davidson, C. Simpson, K. Sandoy, J. Duncan, J. Tate, and

D. Mason. 1979. An ecological study of fishes and

invertebrate macrofauna utilizing the Cape Fear

River Estuary, Carolina Beach Inlet, and adjacent

Atlantic Ocean. Ann. Rep. for 1978. Vol. 15 Instit.

of Marine Sciences, Univ. of North Carolina, Morehead City. NC. 326 p.

Scott, W.B. and M.G. Scott. 1988. Atlantic fishes of

Canada. Can. Bull. Fish. Aquat. Sci., 219. 731 p.
Page 10

Scott, W.B. and S.N. Tibbo. 1968. Food and feeding

habits of swordfish, Xiphias gladius, in the western

North Atlantic. J. Fish. Res. Board Can. 25: 903-919.

Smith, W.G., D.G. McMillan, C. Obenchain, P.

Rosenberg, A. Wells, and M. Silverman. 1980.

Spawning cycles of marine fishes of northeastern

United States based on broadscale surveys of eggs

and larvae, 1977-79. ICES C.M. 1980/L:66. 22 p.

Stender, B.W. and R.M. Martore. 1990. Finfish and

invertebrate communities. In R.F. Van Dolah, P.H.

Wendt, and E.L. Wenner eds. A physical and

ecological characterization of the Charleston Harbor

estuarine system. Chap.VIII, p. 241-288. Mar. Res.

Div., South Carolina Wildl. Mar. Res. Div., Final

Rep. Charleston, SC.

Stillwell, C.E. and N.E. Kohler. 1985. Food and feeding

ecology of the swordfish, Xiphias gladius in the

western North Atlantic Ocean with estimates of daily

ration. Mar. Ecol. Prog. Ser. 22: 239-247.

Stone, S.L., T.A. Lowery, J.D. Field, C.D. Williams,

D.M. Nelson, S.H. Jury, M.E. Monaco, and L.

Andreasen. 1994. Distribution and abundance of

fishes and invertebrates in Mid-Atlantic estuaries.

ELMR Rep. No. 12. NOAA/NOS Strategic Environmental Assessments Division, Silver Spring, MD.

280 p.

Thomas, D.L. and C.B. Milstein. 1973. Ecological studies

in the bays and other waterways near Little Egg Inlet

and in the ocean in the vicinity of the proposed site

for the Atlantic Generating Station, New Jersey.

Progress report for the period January-December

1972. Ichthyological Associates, Inc. Ithaca, NY.

1065 p.

Tibbetts, A.M. 1977. Squid fisheries (Loligo pealei and

Illex illecebrosus) off the northeastern coast of the

United States of America, 1963-1974. Int. Comm.

Northwest Atl. Fish. (ICNAF) Sel. Pap. 2: 85-109.

Wang, J.C.S. and R.J. Kernehan. 1979. Fishes of the

Delaware estuaries: A guide to the early life histories.

EA Communications, Ecological Analysts, Inc.,

Towson, MD. 410 p.

Waring, G. 1975. A preliminary analysis of status of

butterfish in ICNAF Subarea 5 and Statistical Area 6.

Int. Comm. Northwest Atl. Fish. (ICNAF) Res. Doc.,

75/74. 27 p.

Waring, G.T. and S. Murawski. 1982. Butterfish. In M.D.

Grosslein and T.R. Azarovitz eds. Fish distribution.

p. 105-107. MESA New York Bight Atlas Monograph 15. N.Y. Sea Grant Institute. Albany, NY.

Wenner, E.L., W.P. Coon, III, M.H. Shealy, Jr., and P.A.

Sandifer. 1981. Species assemblages, distribution,

and abundance of fishes and decapod crustaceans

from the Winyah Bay estuarine system, S.C. Contrib.

no. 137, South Carolina Mar. Res. Cen. Tech. Rep.

no. 3. South Carolina Sea Grant Consort. 61 p.

Wheatland, S.B. 1956. Pelagic fish eggs and larvae. In

G.A. Riley et al. eds. Oceanography of Long Island

Sound, 1952-1954. p. 234-314. Bull. Bingham

Oceanogr. Collect. 15.

Wilk, S.J., W.W. Morse, D.E. Ralph, and T.R. Azarovitz.

1977. Fishes and associated environmental data

collected in New York Bight, June 1974-June 1975.

NOAA Tech. Rep. NMFS-SSRF-716. 53 p.

Wilk, S.J., W.W. Morse, and L.L. Stehlik. 1990. Annual

cycles of gonad-somatic indices as indicators of

spawning activity for selected species of finfish

collected from the New York Bight. Fish. Bull. (U.S.)

88: 775-786.

Wilk, S.J. and M.J. Silverman. 1976a. Fish and hydrographic collections made by the research vessels

Dolphin and Delaware II during 1968-1972 from

New York to Florida. NOAA Tech. Rep. NMFSSSRF-697. 159 p.

Wilk, S.J. and M.J. Silverman. 1976b. Summer benthic

fish fauna of Sandy Hook Bay, New Jersey. NOAA

Tech. Rep. NMFS-SSRF-698. 16 p.
Page 11

Table 1. Summary of life history and habitat characteristics for butterfish, Peprilus triacanthus. *

Life Stage Geographic Location Habitat Substrate Temperature Salinity

Eggs

(0.68-0.82 mm

diameter)

Cape Sable, Nova Scotia to Florida;

in spring along edge of continental

shelf from Georges Bank to Cape

Hatteras; found progressively closer

to coast from south to north as water

temperatures increase. Commonly

occur in the saline parts of bays and

estuaries from MA to NY and

Chesapeake Bay in spring and

summer.

Surface waters from continental

shelf into estuaries and bays;

collected to about 60 m deep in shelf

waters. Common in high salinity

zone of estuaries and bays from MA

through VA. MARMAP Survey:

collected in surface waters in 10-

1250 m of water.

Literature: 12.8-

22.5oC;

MARMAP

Survey: 6-26oC;

most eggs

collected between

11-17oC

Estuarine to full

seawater; about

25-33 ppt

Larvae

(2.6-16 mm SL)

Cape Sable, Nova Scotia to Cape

Kennedy, FL; most abundant in

central Middle Atlantic Bight in

summer, but absent in the winter.

Commonly occur in bays and

estuaries from MA to NY and

Chesapeake Bay in summer and fall.

Surface waters from continental

shelf into estuaries and bays;

collected to about 60 m deep in shelf

waters; common in high salinity

zone of estuaries and bays; may

spend day deeper in the water

column and migrate to the surface at

night. MARMAP Survey: collected

in surface waters in water 10-1750 m

deep.

Literature: 4.4-

27.9oC.

MARMAP

Survey: 7-26oC;

most eggs

collected between

9-19oC

6.4-37.4 ppt

Juveniles

(16 mm SL120 mm FL)

Cape Sable, Nova Scotia to Florida;

most abundant in Middle Atlantic

Bight in summer and near the edge

of continental shelf in winter.

Commonly occur in bays and

estuaries from MA to VA from

spring through fall; less abundant in

bays and estuaries in the Gulf of

Maine and in the South Atlantic

Bight.

From surface waters to depth on

continental shelf; into coastal bays

and estuaries; common in inshore

areas, including the surf zone, and in

high salinity and mixed salinity

zones of bays and estuaries. NEFSC

Trawl Survey: collected on

continental shelf in 10-330 m of

water; most collected in < 120 m

Larger

individuals

found over

sandy and

muddy

substrates.

4.4-29.7oC;

survival reduced

below 10oC

3.0-37.4 ppt

Adults

(> 120 mm FL)

Cape Sable, Nova Scotia to Florida;

most abundant inshore in Middle

Atlantic Bight in summer and near

the edge of continental shelf in

winter; most abundant north of Cape

Cod in summer and fall; commonly

occur in bays and estuaries from MA

to VA from spring through fall; less

abundant in bays and estuaries in the

Gulf of Maine and in the South

Atlantic Bight; do not migrate far

offshore in South Atlantic Bight.

From surface waters to depths of

270-420 m on continental shelf; into

coastal bays and estuaries; common

in inshore areas, including the surf

zone, and in high salinity and mixed

salinity zones of bays and estuaries.

NEFSC Trawl Survey: collected on

continental shelf in 10-360 m of

water; most collected in < 180 m.

Schools found

over sandy,

sandy-silt, and

muddy

substrates.

4.4-26.0oC;

survival reduced

below 10oC

3.8-33.0 ppt

Spawning

Adults

At least the Gulf of Maine to the

South Atlantic Bight (SAB); most

abundant in Middle Atlantic Bight;

in SAB between Cape Hatteras and

Cape Kennedy. Common in Long

Island Sound, some Long Island

bays, and Chesapeake Bay in spring

and summer. In NY Bight, caught

from May-August.

Spawning occurs on continental

shelf, inshore areas, and in bays and

estuaries (rarely in bays and

estuaries north of Cape Cod).

Spawning adults common in Long

Island Sound and bays and estuaries

of Long Island. In NY Bight, caught

between 3-145 m.

Spawning does not

occur at < 15oC
Page 12

Table 1. cont’d.

Life Stage Dissolved

Oxygen

Light Currents Prey Predators Notes

Eggs

(0.68-0.82 mm

diameter)

Incubation period 2-3 days.

Salinity range based on 78-

100% seawater (Martin and

Drewry, 1978) assuming

seawater at 33 ppt.

Larvae

(2.6-16 mm SL)

More nektonic than

planktonic by 10-15 mm.

Juveniles

(16 mm SL120 mm FL)

HudsonRaritan Bay:

3-9 mg/l; most

5-8 mg/l

Larger juveniles

are pelagic

schoolers; may

congregate near

bottom during

day and disperse

upward at night.

Feed mainly on

planktonic prey,

including thaliaceans,

squids, copepods,

amphipods, decapods,

coelenterates,

polychaetes, small

fishes, and

ctenophores.

Preyed on by

haddock, silver hake,

bluefish, swordfish,

weakfish, goosefish,

sharks, and longfinned squid

Smaller juveniles may

associate with floating objects

including jellyfish and

inanimate objects.

Adults

(> 120 mm FL)

Abundance

declines in

Long Island

Sound at 2.0-

2.9 mg/l.

HudsonRaritan Bay:

3-10 mg/l;

most 6-9 mg/l.

Feed mainly on

planktonic prey,

including thaliaceans,

squids, copepods,

amphipods, decapods,

coelenterates,

polychaetes, small

fishes, and

ctenophores.

Preyed on by

haddock, silver hake,

bluefish, swordfish,

weakfish, goosefish,

sharks, skates, and

long-finned squid

Median size of sexual

maturity 120 mm FL based on

O’Brien et al. (1993).

Spawning

Adults

Spawning occurs JulyOctober on Scotian Shelf,

May-August in Gulf of

Maine, May-October in

Middle Atlantic Bight (peak

June-August), January-April

off Cape Hatteras (peak in

March), and year round in

South Atlantic Bight (peak in

spring).

*In addition to the citations mentioned in the text, the following references were used to compile Table 1: Austin 1973,

1976; Berrien et al. 1978; Colton et al. 1979; Edwards et al. 1962; Lang 1974; Lessard 1974; Obenchain 1981; Wilk

and Silverman 1976a; Wilk et al. 1977.
Page 13

Table 2a. Relative abundance of eggs, larvae, and juvenile butterfish (Peprilus triacanthus) in New England and MidAtlantic estuaries by salinity zone [based on Estuarine Living Marine Resources (ELMR) data in 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, na = no data available.

Eggs Larvae Juveniles

T M S T M S T M S

Passamaquoddy Bay na na na na R R

Englishman/Machias Bays R R

Narraguagus Bay R R

Blue Hill Bay R R

Penobscot Bay R R R R R R

Muscongus Bay R R

Damariscotta River R R

Sheepscot River R R

Kennebec/Androscoggin Rivers R R

Casco Bay R R

Saco Bay R R

Wells Harbor •••

Great Bay R R R R R R

Merrimack River R • R • R •

Massachusetts Bay • • C • • R • • C

Boston Harbor • C • C • R

Cape Cod Bay • C • R • C C

Waquoit Bay R C R C R C

Buzzards Bay R C R C C H

Narragansett Bay R H R C C H

Long Island Sound C C R H A

Connecticut River C

Gardiners Bay C C C

Great South Bay, NY C C R C

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

Barnegat Bay, NJ R C C

New Jersey Inland Bays R C C
Page 14

Table 2a cont’d.

Eggs Larvae Juveniles

T M S T M S T M S

Delaware Bay R C C C

Delaware Inland Bays C

Chincoteague Bay

Chesapeake Bay Mainstream C C C C C C

Chester River

Coptank River R

Patuxent River R

Potomac River R

Tangier/Pocomoke Sound R

Rappahannock River R

York River, VA C

James River, VA C

South Atlantic estuaries – see below

Butterfish occur in estuaries between North Carolina and Florida, but this species was not included in the

ELMR survey of the southeast estuaries (Nelson et al. 1991). Information on their occurrence in South

Atlantic estuaries is presented below.

North Carolina

• Cape Fear River estuary: butterfish < 0.05% of all fishes caught (Schwartz et al. 1979)

South Carolina

• Winyah Bay estuary: butterfish (50-110 mm TL) collected in lower and middle estuary; < 1% of all fishes

caught (Wenner et al. 1981)

• Charleston Harbor estuary system: occur in Charleston Harbor and lower reaches of Ashley, Cooper, and

Wando rivers; < 0.05% of all fishes collected (Stender and Martore 1990)

Georgia

• Sapelo Sound: butterfish collected “occasionally” on ocean beaches and in the lower and middle reaches

of estuary; did not occur at salinities < 19.5 ppt (Dahlberg 1972).

Florida

• Pensacola Bay: juveniles present in winter, spring, summer; rare to uncommon (Cooley 1978).

• Santa Rosa Sound: juveniles collected in winter, spring, summer; rare to uncommon (Cooley 1978).

• Escambia Bay: juveniles collected in winter, spring, fall; rare to uncommon (Cooley 1978).

• Butterfish recorded from ocean beaches on Atlantic and Gulf coasts (to Mississippi) and in Tampa Bay

(Powell et al. 1972).
Page 15

Table 2b. Relative abundance of spawning adult and adult butterfish (Peprilus triacanthus) in New England and MidAtlantic estuaries by salinity zone [based on Estuarine Living Marine Resources (ELMR) data in 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, na = no data available.

Spawning Adults Adults

T M S T M S

Passamaquoddy Bay R R

Englishman/Machias Bays R R

Narraguagus Bay R R

Blue Hill Bay R R

Penobscot Bay R R

Muscongus Bay R R

Damariscotta River R R

Sheepscot River R R

Kennebec/Androscoggin Rivers R R

Casco Bay R R

Saco Bay R R

Wells Harbor • •

Great Bay R R

Merrimack River • • R

Massachusetts Bay • • R • • C

Boston Harbor • • R R

Cape Cod Bay • • C C

Waquoit Bay • R • R C

Buzzards Bay • R • C H

Narragansett Bay R C A

Long Island Sound C A H

Connecticut River • C •

Gardiners Bay • C • C C

Great South Bay, NY • C • R C

Hudson River/Raritan Bay C C

Barnegat Bay, NJ R R

New Jersey Inland Bays R
Page 16

Table 2b cont’d.

Spawning Adults Adults

T M S T M S

Delaware Bay R R C

Delaware Inland Bays • • C

Chincoteague Bay •• ••

Chesapeake Bay Mainstream C C C C

Chester River • •

Coptank River • R •

Patuxent River • R •

Potomac River • R •

Tangier/Pocomoke Sound • •• R •

Rappahannock River • R •

York River, VA • C •

James River, VA • C •

South Atlantic estuaries1

1

See note at bottom of Table 2a.
Page 17

Table 3. Abundance of butterfish eggs, larvae, juveniles, adults, and spawning adults in New England and Mid-Atlantic

estuaries by month summarized across salinity zones [based on Estuarine Living Marine Resources (ELMR) data in

Stone et al. 1994]. Maximum abundance: A = abundant, C = common, R = rare, blank = not present.

Eggs Larvae Juveniles Adults Spawning Adults

Estuary months

present

max.

abun.

months present max.

abun.

months

present

max.

abun.

months

present

max.

abun.

months present max.

abun.

Passamaquoddy Bay ------------ ------------ -----JJASO-- R -----JJASO-- R ------------

Englishman/Machias Bays ------------ ------------ -----JJASO-- R -----JJASO-- R ------------

Narraguagus Bay ------------ ------------ -----JJASO-- R -----JJASO-- R ------------

Blue Hill Bay ------------ ------------ -----JJASO-- R -----JJASO-- R ------------

Penobscot Bay ------JAS--- R ------JAS--- R -----JJASO-- R -----JJASO-- R ------------

Muscongus Bay ------------ ------------ -----JJASO-- R -----JJASO-- R ------------

Damariscotta River ------------ ------------ -----JJASO-- R -----JJASO-- R ------------

Sheepscot River ------------ ------------ -----JJASO-- R -----JJASO-- R ------------

Kennebec/Androscoggin Rivers ------------ ------------ -----JJASO-- R -----JJASO-- R ------------

Casco Bay ------------ ------------ -----JJASO-- R -----JJASO-- R ------------

Saco Bay ------------ ------------ -----JJASO-- R -----JJASO-- R ------------

Wells Harbor ------------ ------------ ------------ ------------ ------------

Great Bay -----JJAS--- R -----JJAS--- R -----JJASO-- R -----JJASO-- R ------------

Merrimack River -----JJA---- R -----JJA---- R -----JJAS--- R -----JJAS--- R ------------

Massachusetts Bay -----JJAS--- C -----JJAS--- R -----JJASO-- C -----JJASO-- C -----JJAS--- R

Boston Harbor -----JJAS--- C ------JAS--- C -----JJASO-- R -----JJASO-- R ------------

Cape Cod Bay -----JJASO-- C ------JASO-- R -----JJASO-- C -----JJASO-- C ------------

Waquoit Bay ----MJJA---- C -----JJASO-- C ----MJJASO-- C ----MJJASO-- C ----MJJAS--- R

Buzzards Bay ----MJJAS--- C -----JJASO-- C ---AMJJASOND A ---AMJJASOND A -----JJAS--- R

Narragansett Bay ----MJJA---- A -----JJASO-- C ---AMJJASOND A ---AMJJASOND A ----MJJA---- R

Gardiners Bay ----MJJ----- C ----MJJ----- C ----MJJASOND C ----MJJASOND C ----MJJ----- C

Long Island Sound -----JJAS--- C -----JJASON- C ----MJJASOND A ----MJJASOND A -----JJAS--- C

Connecticut River ------------ ------------ ----MJJASOND C ----MJJASOND C ------------

Great South Bay ----MJJ----- C ----MJJA---- C ----MJJASOND C ----MJJASOND C ----MJJ----- C

Hudson River/Raritan Bay -----JJA---- R ----MJJASON- C ---AMJJASON- C ---AMJJASON- C ------------

Barnegat Bay ------------ -----JJA---- R -----JJASO-- C ----MJJASO-- R ------------

New Jersey Inland Bays ------------ -----JJA---- R -----JJASO-- C ------JAS--- R ------------

Delaware Bay ----MJJ----- R ----MJJ----- C ------JASOND C ----MJJASO-- C ----MJJ----- R

Delaware Inland Bays ------------ ------------ ----MJJASON- C ----MJJASON- C ------------

Chincoteague Bay ------------ ------------ ------------ ------------ ------------

Chesapeake Bay ----MJJ----- C -----JJA---- C ------JASO-- C ---AMJJASON- C ----MJJ----- C

Potomac River ------------ ------------ -----JJASO-- R ----MJJASO-- R ------------

Rappahannock River ------------ ------------ ------JASON- R ---AMJJASON- R ------------

York River ------------ ------------ ------JASON- C ---AMJJASON- C ------------

James River ------------ ------------ ------JASON- C ---AMJJASON- C ------------

Patuxent River ------------ ------------ ------JAS--- R -----JJAS--- R ------------

Chester River ------------ ------------ ------------ ------------ ------------

Choptank River ------------ ------------ ------JAS--- R -----JJAS--- R ------------

Tangier/Pocomoke Sound ------------ ------------ ------JASO-- R ----MJJASO-- R ------------
Page 18

Figure 1. The adult butterfish, Peprilus triacanthus (from Goode 1884).
Page 19

Figure 2. The distribution of butterfish from Newfoundland to Cape Hatteras. Data are from the U.S. NOAA/Canada

DFO East Coast of North America Strategic Assessment Project (http://www-orca.nos.noaa.gov/projects/ecnasap/

ecnasap_table1.html).
Page 20

Figure 3. Abundance (percent occurrence) of the major prey items of butterfish collected during NEFSC bottom trawl

surveys from 1973-1980 and 1981-1990. The 1-10 cm size range corresponds, at least roughly, to juveniles, and the 11-

30 cm size class corresponds to adults. The category “animal remains” refers to unidentifiable animal matter. Methods

for sampling, processing, and analysis of samples differed between the time periods [see Reid et al. (1999) for details].

Unknown Animal Remains 52.9%

Arthropoda 21.7%

Urochordata 13.2%

Cnidaria 3.3%

Annelida 2.2%

Chaetognatha 2.2%

Mollusca 2.2%

All Other Groups 2.2%

Unknown Animal Remains 43.5%

Arthropoda 24.9%

Urochordata 8.6%

Platyhelminthes 6.5%

Annelida 5.2%

Mollusca 3.8%

Chaetognatha 1.8%

Cnidaria 1.4%

All Other Groups 4.3%

Unknown Animal Remains 100.0%

Unknown Animal Remains 76.3%

Arthropoda 9.7%

Plankton 6.5%

Annelida 3.2%

Fish 3.2%

Mollusca 1.1%

1-10 cm

(n = 469)

1-10 cm

(n = 9)

11-30 cm

(n = 893)

11-30 cm

(n = 74)

a) 1973-1980

b) 1981-1990
Page 21

Figure 4. Abundance of butterfish eggs relative to water column temperature (to a maximum of 200 m) and bottom

depth from NEFSC MARMAP ichthyoplankton surveys (1978-1987) by month for all years combined. Open bars

represent the proportion of all stations surveyed, while solid bars represent the proportion of the sum of all standardized

catches (number/10 m2

).

April

0

10

20

30

40

50

Stations

Egg Catch

May

0

10

20

30

June

Percent

0

10

20

30

40

July

0

10

20

30

August

0

10

40

50

Butterfish Eggs

September

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

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

0

10

20

50

60

April

0

10

20

30

40

50

60

Stations

Egg Catch

Butterfish Eggs

September

Bottom Depth (m), Interval Midpoint

10

30

50

70

90

110

130

150

170

190

210

230

250

270

290

325

375

450

750

1250

1750

>2000

0

10

20

30

60

70

May

0

10

20

June

Percent

0

10

20

30

July

0

10

20

30

August

0

10

20

30

40

50
Page 22

Figure 5. Abundance of butterfish larvae (< 14 mm) relative to water column temperature (to a maximum of 200 m) and

bottom depth from NEFSC MARMAP ichthyoplankton surveys (1977-1987) by month for all years combined. Open

bars represent the proportion of all stations surveyed, while solid bars represent the proportion of the sum of all

standardized catches (number/10 m2

).

January

0

10

20

30

90

100

Stations

Larva Catch

April

0

10

20

30

May

0

10

20

June

Percent

0

10

20

30

40

July

0

10

20

30

August

0

10

20

November

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

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

0

20

40

60

Butterfish Larvae (<14.0 mm Length)

September

0

10

20

30

March

0

10

20

30

40

October

0

10

20

February

0

20

40

60

January

0

10

20

90

100

Stations

Larva Catch

April

0

10

20

30

40

Butterfish Larvae (<14.0 mm Length)

September

0

10

20

30

40

May

0

10

20

30

June

Percent

0

10

20

30

40

July

0

10

20

30

40

50

August

0

10

20

30

40

October

0

10

20

30

March

0

10

20

30

70

80

February

0

20

40

60

November

Bottom Depth (m), Interval Midpoint

10

30

50

70

90

110

130

150

170

190

210

230

250

270

290

325

375

450

750

1250

1750

>2000

0

10

20

30

40

60

70
Page 23

Figure 6. Abundance of juvenile (< 12 cm) and adult (≥ 12 cm) butterfish relative to bottom water temperature and

depth based on NEFSC bottom trawl surveys (1963-1997) by season for all years combined. Open bars represent the

proportion of all stations surveyed, while solid bars represent the proportion of the sum of all standardized catches

(number/10 m2

).

SPRING

0

5

10

15

20

25

30

4681

0

1

2

1

4

1

6

1

8

2

0

BOTTOM TEMPERATURE (C)

PERCENT

STATIONS N=904

CATCHES N=145432

SUMMER

0

5

10

15

20

25

30

5791

1

1

3

1

5

1

7

1

9

2

1

2

3

2

5

2

7

BOTTOM TEMPERATURE (C)

PERCENT

STATIONS N=409

CATCHES N=557961

FALL

0

5

10

15

20

25

30

5791

1

1

3

1

5

1

7

1

9

2

1

2

3

2

5

2

7

BOTTOM TEMPERATURE (C)

PERCENT

STATIONS N=3258

CATCHES N=1081500

WINTER

0

5

10

15

20

25

30

4567891

0

1

1

1

2

1

3

BOTTOM TEMPERATURE (C)

PERCENT

STATIONS N=259

CATCHES N=48506

Juveniles: < 12 cm TL

SPRING

0

10

20

30

40

50

60

70

0 3

0

6

0

9

0

1

2

0

1

5

0

1

8

0

2

1

0

2

4

0

2

7

0

3

1

0

3

6

0

DEPTH (m)

PERCENT

STATIONS N=1057

CATCHES N=161518

SUMMER

0

10

20

30

40

50

60

70

0 2

0

4

0

6

0

8

0

1

0

0

1

3

0

1

5

0

2

1

0

DEPTH (m)

PERCENT

STATIONS N=447

CATCHES N=567902

FALL

0

10

20

30

40

50

60

70

0 3

0

6

0

9

0

1

2

0

1

5

0

1

8

0

2

1

0

2

4

0

2

7

0

3

0

0

3

3

0

3

8

0

DEPTH (m)

PERCENT

STATIONS N=3939

CATCHES N=1337946

WINTER

0

10

20

30

40

50

60

70

2

0

4

0

6

0

8

0

1

0

0

1

2

0

1

4

0

1

6

0

1

9

0

2

1

0

3

1

0

DEPTH (m)

PERCENT

STATIONS N=267

CATCHES N=48737
Page 24

Figure 6. cont’d.

SPRING

0

5

10

15

20

25

30

4681

0

1

2

1

4

1

6

1

8

2

0

BOTTOM TEMPERATURE (C)

PERCENT

STATIONS N=1480

CATCHES N=244695

SUMMER

0

5

10

15

20

25

30

25791

1

1

3

1

5

1

7

1

9

2

1

2

3

2

5

2

7

BOTTOM TEMPERATURE (C)

PERCENT

STATIONS N=486

CATCHES N=106730

FALL

0

5

10

15

20

25

30

4681

0

1

2

1

4

1

6

1

8

2

0

2

2

2

4

2

6

BOTTOM TEMPERATURE (C)

PERCENT

STATIONS N=3472

CATCHES N=711068

WINTER

0

5

10

15

20

25

30

34567891

0

1

1

1

2

1

3

BOTTOM TEMPERATURE (C)

PERCENT

STATIONS N=349

CATCHES N=45058

Adults: ≥ 12 cm TL

SPRING

0

5

10

15

20

25

30

0 3

0

6

0

9

0

1

2

0

1

5

0

1

8

0

2

1

0

2

4

0

2

7

0

3

0

0

3

3

0

3

6

0

4

9

0

DEPTH (m)

PERCENT

STATIONS N=1705

CATCHES N=264295

SUMMER

0

5

10

15

20

25

30

0 2

0

4

0

6

0

8

0

1

0

0

1

2

0

1

4

0

1

6

0

2

4

0

DEPTH (m)

PERCENT

STATIONS N=513

CATCHES N=109454

FALL

0

5

10

15

20

25

30

0 3

0

6

0

9

0

1

2

0

1

5

0

1

8

0

2

1

0

2

4

0

2

7

0

3

0

0

3

3

0

3

8

0

DEPTH (m)

PERCENT

STATIONS N=4116

CATCHES N=891263

WINTER

0

5

10

15

20

25

30

2

0

4

0

6

0

8

0

1

0

0

1

2

0

1

4

0

1

6

0

1

8

0

2

0

0

2

2

0

2

4

0

2

7

0

3

1

0

DEPTH (m)

PERCENT

STATIONS N=359

CATCHES N=45303
Page 25

Figure 7. Abundance of juvenile and adult butterfish relative to bottom water temperature and depth based on

Massachusetts inshore bottom trawl surveys (spring and autumn 1978-1996, all years combined). Open bars represent

the proportion of all stations surveyed, while solid bars represent the proportion of the sum of all standardized catches

(number/10 m2

).

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

0

10

20

30

40

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

0

5

10

15

20

25

0

20

40

60

0

10

20

30

40

0

5

10

15

20

25

0

10

20

30

40

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

0

4

8

12

16

20

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

0

10

20

30

40

Juveniles Adults

Stations

Catches

Spring Spring

Spring Spring

Autumn Autumn

Autumn Autumn

Bottom Depth (m) Bottom Depth (m)

Bottom Depth (m) Bottom Depth (m)

Bottom Temperature (C) Bottom Temperature (C)

Bottom Temperature (C) Bottom Temperature (C)

Mass. Inshore Trawl Surveys

Butterfish
Page 26

Figure 8. Abundance of juvenile and adult butterfish relative to bottom water temperature, depth, dissolved oxygen and

salinity from Hudson-Raritan estuary trawl surveys (1992-1997) for all years combined.

Temperature (C)

Depth (ft)

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

0

5

10

15

20

25

Stations

Catches

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

0

5

10

15

20

25

30

35

Dissolved Oxygen (mg/l)

Salinity (ppt)

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

0

5

10

15

20

25

30

35

40

45

50

15 17 19 21 23 25 27 29 31 33 35

0

2

4

6

8

10

12

14

16

18

Juveniles ( 12 cm) <

Temperature (C)

Depth (ft)

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

0

2

4

6

8

10

12

14

16

Stations

Catches

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

0

5

10

15

20

25

30

35

Dissolved Oxygen (mg/l)

Salinity (ppt)

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

0

5

10

15

20

25

30

35

15 17 19 21 23 25 27 29 31 33 35

0

2

4

6

8

10

12

14

16

Adults ( 12 cm) >
Page 27

Figure 9. Distribution of butterfish eggs based on NEFSC MARMAP ichthyoplankton surveys from April to September,

1978-1987 [see Reid et al. (1999) for details].

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

35

36

37

38

39

40

41

42

43

44

45

Butterfish

Eggs

MARMAP Ichthyoplankton Surveys

61-cm Bongo Net; 0.505-mm mesh

April to September; 1978 to 1987

Number of tows = 5205; with eggs = 688

Eggs / 10m2

1 to <10

10 to <100

100 to <1000

1000 to <10000

10000 to 13222

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

35

36

37

38

39

40

41

42

43

44

45

Butterfish

Eggs

MARMAP Ichthyoplankton Surveys

61-cm Bongo Net; 0.505-mm mesh

April; 1978 to 1987

Number of tows = 1020; with eggs = 10

None

1 to <10

10 to <100

100 to 129

Eggs / 10m2

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

35

36

37

38

39

40

41

42

43

44

45

Butterfish

Eggs

MARMAP Ichthyoplankton Surveys

61-cm Bongo Net; 0.505-mm mesh

May; 1978 to 1987

Number of tows = 1085; with eggs = 167

None

1 to <10

10 to <100

100 to <1000

Eggs / 10m2

1000 to 4843

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

35

36

37

38

39

40

41

42

43

44

45

Butterfish

Eggs

MARMAP Ichthyoplankton Surveys

61-cm Bongo Net; 0.505-mm mesh

June; 1978 to 1987

Number of tows = 709; with eggs = 123

None

1 to <10

10 to <100

100 to <1000

Eggs / 10m2

1000 to 5892
Page 28

Figure 9. cont’d.

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

35

36

37

38

39

40

41

42

43

44

45

Butterfish

Eggs

MARMAP Ichthyoplankton Surveys

61-cm Bongo Net; 0.505-mm mesh

July; 1978 to 1987

Number of tows = 781; with eggs = 235

None

1 to <10

10 to <100

100 to <1000

Eggs / 10m2

1000 to 1652

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

35

36

37

38

39

40

41

42

43

44

45

Butterfish

Eggs

MARMAP Ichthyoplankton Surveys

61-cm Bongo Net; 0.505-mm mesh

August; 1978 to 1987

Number of tows = 863; with eggs = 141

None

1 to <10

10 to <100

100 to <1000

Eggs / 10m2

1000 to <10000

10000 to 13222

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

35

36

37

38

39

40

41

42

43

44

45

Butterfish

Eggs

MARMAP Ichthyoplankton Surveys

61-cm Bongo Net; 0.505-mm mesh

September; 1978 to 1987

Number of tows = 747; with eggs = 12

None

1 to <10

10 to <100

100 to 211

Eggs / 10m2
Page 29

Figure 10. Distribution of butterfish larvae (< 14 mm) collected during NEFSC MARMAP ichthyoplankton surveys

from January through November, 1977-1987 [see Reid et al. (1999) for details].

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

35

36

37

38

39

40

41

42

43

44

45

MARMAP Ichthyoplankton Surveys

61-cm Bongo Net; 0.505-mm mesh

January to November (1977 to 1987)

Butterfish

Number of Tows = 10835, with larvae = 1053

larvae <14.0mm

1 to < 10

10 to < 100

100 to < 1000

1000 to 1765

Number of 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

January (1977 to 1987)

Number of Tows = 434, with larvae = 1

Number of Larvae / 10m2

None

1 to 3

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

35

36

37

38

39

40

41

42

43

44

45

February (1977 to 1987)

Number of Tows = 686, with larvae = 2

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

March (1977 to 1987)

Number of Tows = 1031, with larvae = 3

Number of Larvae / 10m2

None

1 to < 10

10 to 19
Page 30

Figure 10. cont’d.

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

35

36

37

38

39

40

41

42

43

44

45

April (1977 to 1987)

Number of Tows = 1281, with larvae = 21

Number of Larvae / 10m2

None

1 to < 10

10 to 59

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

35

36

37

38

39

40

41

42

43

44

45

May (1977 to 1987)

Number of Tows = 1472, with larvae = 67

Number of Larvae / 10m2

None

1 to < 10

10 to < 100

100 to 391

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

35

36

37

38

39

40

41

42

43

44

45

June (1977 to 1987)

Number of Tows = 893, with larvae = 101

None

1 to < 10

10 to < 100

100 to < 1000

1000 to 1365

Number of 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

July (1977 to 1987)

Number of Tows = 938, with larvae = 343

None

1 to < 10

10 to < 100

100 to < 1000

1000 to 1765

Number of Larvae / 10m2
Page 31

Figure 10. cont’d.

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

35

36

37

38

39

40

41

42

43

44

45

August (1977 to 1987)

Number of Tows = 1148, with larvae = 408

Number of Larvae / 10m2

None

1 to < 10

10 to < 100

100 to 477

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

35

36

37

38

39

40

41

42

43

44

45

September (1977 to 1987)

Number of Tows = 774, with larvae = 69

Number of Larvae / 10m2

None

1 to < 10

10 to < 100

100 to 108

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

35

36

37

38

39

40

41

42

43

44

45

October (1977 to 1987)

Number of Tows = 1147, with larvae = 35

Number of Larvae / 10m2

None

1 to < 10

10 to 45

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

35

36

37

38

39

40

41

42

43

44

45

November (1977 to 1987)

Number of Tows = 1031, with larvae = 3

Number of Larvae / 10m2

None

1 to 5
Page 32

Figure 11. Distribution of juvenile and adult butterfish collected during NEFSC bottom trawl surveys during all seasons

during 1963-1997. Densities are represented by dot size in spring and fall plots, while only presence and absence are

represented in winter and summer plots [see Reid et al. (1999) for details].
Page 33

Figure 11. cont’d.
Page 34

Figure 12. Distribution of juvenile and adult butterfish in Massachusetts coastal waters during spring and autumn

Massachusetts trawl surveys, 1978-1996 [see Reid et al. (1999) for details].

Butterfish

Mass. Inshore Trawl Survey

 Spring 1978 - 1996

 Juveniles (<12cm)

Number/Tow

 1 to <10

 10 to <25

 25 to <50

 50 to <100

 100 to <226

Butterfish

Mass. Inshore Trawl Survey

 Autumn 1978 - 1996

 Juveniles (<12cm)

Number/Tow

 1 to <100

 100 to <500

 500 to <1000

 1000 to <5000

 5000 to <18170

Butterfish

Mass. Inshore Trawl Survey

 Spring 1978 - 1996

 Adults (>=12cm)

Number/Tow

 1 to <25

 25 to <50

 50 to <100

 100 to <500

 500 to <1008

Butterfish

Mass. Inshore Trawl Survey

 Autumn 1978 - 1996

 Adults (>=12cm)

Number/Tow

 1 to <50

 50 to <100

 100 to <500

 500 to <1000

 1000 to <1585
Page 35

Figure 13. Distribution of juvenile and adult butterfish in the SEAMAP bottom trawl surveys in all seasons for all years

combined (1986-1996).

Number/Tow

 0 to <0

 1 to <5

 5 to <7

 7 to <10

 10 to <12

 12 to <15

Butterfish

SEAMAP 1986 - 1996

Winter Juveniles (<12cm)

Number/Tow

 0 to <0

 1 to <25

 25 to <50

 50 to <100

 100 to <200

 200 to <291

Butterfish

SEAMAP 1986 - 1996

Spring Juveniles (<12cm)

Number/Tow

 0 to <0

 1 to <25

 25 to <50

 50 to <100

 100 to <200

 200 to <471

Butterfish

SEAMAP 1986 - 1996

Summer Juveniles (<12cm)

Number/Tow

 0 to <0

 1 to <25

 25 to <50

 50 to <100

 100 to <150

 150 to <156

Butterfish

SEAMAP 1986 - 1996

Autumn Juveniles (<12cm)
Page 36

Figure 13. cont’d.

Number/Tow

 0 to <0

 1 to <25

 25 to <50

 50 to <100

 100 to <150

 150 to <177

Butterfish

SEAMAP 1986 - 1996

Winter Adults (>=12cm)

Number/Tow

 0 to <0

 1 to <25

 25 to <50

 50 to <100

 100 to <200

 200 to <351

Butterfish

SEAMAP 1986 - 1996

Spring Adults (>=12cm)

Number/Tow

 0 to <0

 1 to <10

 10 to <25

 25 to <50

 50 to <100

 100 to <111

Butterfish

SEAMAP 1986 - 1996

Summer Adults (>=12cm)

Number/Tow

 0 to <0

 1 to <25

 25 to <50

 50 to <100

 100 to <200

 200 to <297

Butterfish

SEAMAP 1986 - 1996

Autumn Adults (>=12cm)
Page 37

Figure 14. Distribution of juvenile and adult butterfish collected in Narragansett Bay during 1990-1996 Rhode Island

bottom trawl surveys. The numbers shown at each station are the average catch per tow rounded to one decimal place

[see Reid et al. (1999) for details].

0.3

0.4

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.1

0.0 0.0

0.1

0.0

0.1

0.0

0.0

0.0

0.0

0.2

0.1

0.3

0.2

0.2 0.0

109.8

83.8

78.3

17.5

155.5

71.2

89.5

219.4

44.5

221.4

371.0

319.4 228.9

586.1

45.3

36.4

2.8

124.6

8.0

1.0

45.4

22.7

187.6

174.2

62.8 34.4

Spring

Summer Autumn

Winter

Butterfish Juveniles (< 12 cm)
Page 38

Figure 14. cont’d.

1.2

0.5

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.4

0.0 0.0

0.9

0.0

0.0

0.0

0.0

0.0

0.0

1.9

25.3

33.2

5.3

2.8 0.0

261.8

3.6

4.0

0.9

6.3

1.1

0.4

14.8

15.6

48.1

73.3

68.4 1.9

200.8

32.4

17.0

3.9

112.5

2.9

0.1

17.8

6.7

130.8

33.8

2.4 22.5

Spring

Summer Autumn

Winter

Butterfish Adults (>= 12 cm)
Page 39

Figure 15. Distribution of juvenile and adult butterfish in Long Island Sound in spring and autumn, from the

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

Number/Tow

 1 to <25

 25 to <50

 50 to <100

 100 to <500

 500 to <1218

Number/Tow

 1 to <500

 500 to <1000

 1000 to <2000

 3000 to <4000

 4000 to <10200

State of Connecticut DEP

Fisheries Division

Long Island Sound Trawl Survey

SPRING 1992 - 1997

State of Connecticut DEP

Fisheries Division

Long Island Sound Trawl Survey

AUTUMN 1992 - 1997

Butterfish

Butterfish
Page 40

Figure 16. Distribution of juvenile and adult butterfish in the Hudson-Raritan estuary based on Hudson-Raritan trawl

surveys, 1992-1997 [see Reid et al. (1999) for details].

Staten

Island

NEW

YORK

NEW

JERSEY

No/Tow

 1 - 9

10 - 24

25 - 49

50 - 99

100 - 240

Butterfish

Juveniles ( 12 cm) <

Hudson-Raritan Estuary

Spring 1992 - 1997

Staten

Island

NEW

YORK

NEW

JERSEY

No/Tow

 1 - 9

10 - 24

25 - 49

50 - 99

100 - 240

Butterfish

Juveniles ( 12 cm) <

Hudson-Raritan Estuary

Summer 1992 - 1996

Staten

Island

NEW

YORK

NEW

JERSEY

No/Tow

 1 - 9

10 - 24

25 - 49

50 - 99

100 - 240

Butterfish

Juveniles ( 12 cm) <

Hudson-Raritan Estuary

Fall 1992 - 1996

Staten

Island

NEW

YORK

NEW

JERSEY

No/Tow

 1 - 9

10 - 24

25 - 49

50 - 99

100 - 240

Butterfish

No Catches

<

Hudson-Raritan Estuary

 Winter 1992 - 1997

Juveniles ( 12 cm)
Page 41

Figure 16. cont’d.

Staten

Island

NEW

YORK

NEW

JERSEY

No/Tow

 1 - 9

10 - 24

25 - 49

50 - 99

100 - 240

Butterfish

Hudson-Raritan Estuary

Spring 1992 - 1997

Adults ( 12 cm) >

Staten

Island

NEW

YORK

NEW

JERSEY

No/Tow

 1 - 9

10 - 24

25 - 49

50 - 99

100 - 240

Butterfish

Adults ( 12 cm) >

Hudson-Raritan Estuary

Fall 1992 - 1996

Staten

Island

NEW

YORK

NEW

JERSEY

No/Tow

 1 - 9

10 - 24

25 - 49

50 - 99

100 - 240

Butterfish

Adults ( 12 cm) >

Hudson-Raritan Estuary

Winter 1992 - 1997

No Catches

Staten

Island

NEW

YORK

NEW

JERSEY

No/Tow

 1 - 9

10 - 24

25 - 49

50 - 99

100 - 240

Hudson-Raritan Estuary

Summer 1992 - 1997

Adults ( 12 cm) >

Butterfish
Page 42

Figure 17. Commercial landings and abundance indices (from the NEFSC bottom trawl surveys) for butterfish from the

Gulf of Maine to the Middle Atlantic.

Gulf of Maine - Middle Atlantic

Year

1960 1965 1970 1975 1980 1985 1990 1995 2000

Landings (mt x 1000)

0

5

10

15

20

25

Stratified mean catch/tow (kg)

0

5

10

15

Landings (mt)

Survey index (kg)

Smoothed survey index (kg)
NORTHEAST FISHERIES SCIENCE CENTER

Dr. Michael P. Sissenwine, Science & Research Director

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

Teri L. Frady, Research Communications Unit Chief

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

Publishing in NOAA Technical Memorandum NMFS-NE

Manuscript Qualification

This series represents a secondary level of scientific publishing in the National Marine Fisheries Service (NMFS). 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 publishing

should be initially submitted to one of NMFSs primary series

(i.e., Fishery Bulletin, NOAA Technical Report NMFS, or Marine Fisheries Review).

Identical, or fundamentally identical, manuscripts should

not be concurrently submitted to this and any other publication

series. Manuscripts which have been rejected by any primary

series strictly because of geographic or temporal limitations may

be submitted to this series.

Manuscripts by Northeast Fisheries Science Center

(NEFSC) authors will be published in this series upon approval

by the NEFSC's Deputy Science & Research Director. Manuscripts by non-NEFSC authors may be published in this series if:

1) the manuscript serves the NEFSCs mission; 2) the manuscript meets the Deputy Science & Research Directors approval; and 3) the author arranges for the printing and binding

funds to be transferred to the NEFSCs Research Communications Unit account from another federal account. For all manuscripts submitted by non-NEFSC authors and published in this

series, the NEFSC will disavow all responsibility for the manuscripts contents; authors must accept such responsibility.

The ethics of scientific research and scientific publishing

are a serious matter. All manuscripts submitted to this series are

expected to adhere -- at a minimum -- to the ethical guidelines

contained in Chapter 1 (Ethical Conduct in Authorship and

Publication) of the CBE Style Manual, fifth edition (Chicago,

IL: Council of Biology Editors). Copies of the manual are

available at virtually all scientific libraries.

Manuscript Preparation

Organization: Manuscripts must have an abstract, table of

contents, and -- if applicable -- lists of tables, figures, and

acronyms. As much as possible, use traditional scientific manuscript organization for sections: Introduction, Study Area,

Methods & Materials, Results, Discussion and/or Conclusions, Acknowledgments, and References Cited.

Style: All NEFSC publication and report series are obligated to conform to the style contained in the most recent edition

of the United States Government Printing Office Style Manual.

That style manual is silent on many aspects of scientific manuscripts. NEFSC publication and report series rely more on the

CBE Style Manual, fifth edition.

For in-text citations, use the name-date system. A special

effort should be made to ensure that the list of cited works

contains all necessary bibliographic information. For abbreviating serial titles in such lists, use the most recent edition of the

Serial Sources for the BIOSIS Previews Database (Philadelphia,

PA: Biosciences Information Service). Personal communications must include date of contact and full name and mailing

address of source.

For spelling of scientific and common names of fishes,

mollusks, and decapod crustaceans from the United States and

Canada, use Special Publications No. 20 (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 Mammalogy (Lawrence, KS). For spelling in

general, use the most recent edition of Websters Third New

International Dictionary of the English Language Unabridged

(Springfield, MA: G.&C. Merriam).

Typing text, tables, and figure captions: Text, including

tables and figure captions, must be converted to, or able to be

coverted to, WordPerfect. In general, keep text simple (e.g.,

dont switch fonts, dont use hard returns within paragraphs,

dont indent except to begin paragraphs). Especially, dont use

WordPerfect graphics for embedding tables and figures in text.

If the automatic footnoting function is used, also save a list of

footnotes as a separate WordPerfect file. When the final draft is

ready for review, save the text, tables, figure captions, footnotes,

and front matter as separate document files.

Tables should be prepared using all tabs or all spaces

between columnar data, but not a combination of the two.

Figures must be original (even if oversized) and on paper; they

cannot be photocopies (e.g., Xerox) unless that is all that is

available, nor be on disk. Except under extraordinary circumstances, color will not be used in illustrations.

Manuscript Submission

Authors must submit one paper copy of the double-spaced

manuscript, one magnetic copy on a disk, and original figures (if

applicable). NEFSC authors must include a completely signedoff NEFSC Manuscript/Abstract/Webpage Review Form.

Non-NEFSC authors who are not federal employees will be

required to sign a Release of Copyright form.

Send all materials and address all correspondence to:

Jon A. Gibson, Biological Sciences Editor

Northeast Fisheries Science Center

National Marine Fisheries Service

166 Water Street

Woods Hole, MA 02543-1026 USA
Research Communications Unit

Northeast Fisheries Science Center

National Marine Fisheries Service, NOAA

166 Water St.

Woods Hole, MA 02543-1026

NOAA Technical Memorandum NMFS-NE -- This series is issued irregularly. The series includes: data reports of longterm or large area studies; synthesis reports for major resources or habitats; annual reports of assessment or monitoring programs;

documentary reports of oceanographic conditions or phenomena; manuals describing field and lab techniques; literature surveys of major

resource or habitat topics; findings of task forces or working groups; summary reports of scientific or technical workshops; and indexed

and/or annotated bibliographies. All issues receive internal scientific review and most issues receive technical and copy editing. Limited

free copies are available from authors or the NEFSC. Issues are also available from the National Technical Information Service, 5285

Port Royal Rd., Springfield, VA 22161.

Fishermen's Report and The Shark Tagger -- The Fishermen's Report (FR) is a quick-turnaround report on the

distribution and relative abundance of commercial fisheries resources as derived from each of the NEFSC's periodic research vessel

surveys of the Northeast's continental shelf. There is no scientific review, nor any technical or copy editing, of the FR; copies are available

through free subscription. The Shark Tagger (TST) is an annual summary of tagging and recapture data on large pelagic sharks as derived

from the NMFS's Cooperative Shark Tagging Program; it also presents information on the biology (movement, growth, reproduction, etc.)

of these sharks as subsequently derived from the tagging and recapture data. There is internal scientific review, but no technical or copy

editing, of the TST; copies are available only to participants in the tagging program.

Northeast Fisheries Science Center Reference Document -- This series is issued irregularly. The series

includes: data reports on field and lab observations or experiments; progress reports on continuing experiments, monitoring, and

assessments; background papers for scientific or technical workshops; and simple bibliographies. Issues receive internal scientific review

but no technical or copy editing. No subscriptions. Free distribution of single copies.

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

benefit of the nation through their science-based conservation and management and promotion of the health of their

environment." As the research arm of the NMFS's Northeast Region, the Northeast Fisheries Science Center (NEFSC)

supports the NMFS mission by "planning, developing, and managing multidisciplinary programs of basic and applied

research to: 1) better understand the living marine resources (including marine mammals) of the Northwest Atlantic, and

the environmental quality essential for their existence and continued productivity; and 2) describe and provide to

management, industry, and the public, options for the utilization and conservation of living marine resources and

maintenance of environmental quality which are consistent with national and regional goals and needs, and with

international commitments." Results of NEFSC research are largely reported in primary scientific media (e.g.,

anonymously-peer-reviewed scientific journals). However, to assist itself in providing data, information, and advice to

its constituents, the NEFSC occasionally releases its results in its own media. Those media are in three categories:

Publications and Reports

of the

Northeast Fisheries Science Center

To obtain a copy of a technical memorandum or a reference document, or to subscribe to the fishermen's report,

write: Research Communications Unit, Northeast Fisheries Science Center, 166 Water St., Woods Hole, MA

02543-1026. An annual list of NEFSC publications and reports is available upon request at the above address.

Any use of trade names in any NEFSC publication or report does not imply endorsement.

STANDARD

 MAIL A