NOAA Technical Memorandum NMFS-NE-137

Essential Fish Habitat Source Document:

Windowpane, Scophthalmus aquosus,

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:

Windowpane, Scophthalmus aquosus,

Life History and Habitat Characteristics

 Sukwoo Chang, Peter L. Berrien, Donna L. Johnson,

and Wallace W. Morse

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

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

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

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

All issues may be cited in formal scientific communications.

NOAA Technical Memorandum NMFS-NE-137
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 ..............................................................................................................................................................................2

Geographical Distribution.........................................................................................................................................................................3

Status of the Stocks ...................................................................................................................................................................................4

Research Needs .........................................................................................................................................................................................4

Acknowledgments.....................................................................................................................................................................................5

References Cited .......................................................................................................................................................................................5

Tables

Table 1. Summary of the distribution and abundance of windowpane in North Atlantic and Mid-Atlantic estuaries .............................8

Table 2. Summary of life history and habitat parameters for windowpane ..............................................................................................9

Figures

Figure 1. The windowpane, Scophthalmus aquosus (from Bigelow and Schroeder 1953)...................................................................10

Figure 2. Distribution and relative abundance of juvenile windowpane from NEFSC bottom trawl surveys.......................................11

Figure 3. Distribution and relative abundance of adult windowpane from NEFSC bottom trawl surveys ...........................................12

Figure 4. Abundance (percent occurrence) of the major prey items of windowpane collected during NEFSC trawl surveys..............13

Figure 5. Percentage of windowpane eggs in relation to water temperature and depth from MARMAP ichthyoplankton surveys......14

Figure 6. Percentage of windowpane larvae in relation to water temperature and depth from MARMAP ichthyoplankton surveys ...15

Figure 7. Percentage of juvenile and adult windowpane in relation to water temperature and depth based on NEFSC surveys..........16

Figure 8. Distribution and relative abundance of juvenile and adult windowpane from Massachusetts inshore trawl surveys............17

Figure 9. Percentage of juvenile and adult windowpane in relation to water temperature and depth from Massachusetts surveys......18

Figure 10. Distribution and relative abundance of juvenile and adult windowpane from the Hudson-Raritan estuary..........................19

Figure 11. Percent frequency of windowpane in relation to bottom temperature, depth, DO, and salinity in the Hudson-Raritan ........21

Figure 12. Abundance and length frequency distributions of windowpane in Long Island Sound from Connecticut trawl surveys......22

Figure 13. Distribution and abundance of windowpane from Newfoundland to Cape Hatteras during 1975-1994 ...............................23

Figure 14. Distribution and abundance of windowpane eggs collected during MARMAP ichthyoplankton surveys ............................24

Figure 15. Distribution and abundance of windowpane larvae collected during MARMAP ichthyoplankton surveys..........................27

Figure 16. Commercial landings and NEFSC bottom trawl indices for windowpane from Gulf of Maine to Middle Atlantic Bight ....31

Figure 17. Distribution and abundance of juveniles and adults during low and high abundance periods from NEFSC surveys ..........32
Page 1

INTRODUCTION

The windowpane, Scophthalmus aquosus, is an

eurythemal, euryhaline, and fast-growing fish with a thin

body (Figure 1). It inhabits estuaries, near-shore waters,

and the continental shelf in the northwest Atlantic.

Windowpane is not a target of the commercial fishing

industry, but is mainly caught as bycatch in bottom trawl

fisheries. It is managed by the New England Fishery

Management Council under the Multispecies Fishery

Management Plan (NEFMC 1993). This Essential Fish

Habitat source document provides information on the life

history and habitat characteristics of windowpane.

LIFE HISTORY

The windowpane is a left-eyed flounder with a thin

body and nearly round outline. It occurs from the Gulf of

Saint Lawrence to Florida (Scott and Scott 1988), but is

most abundant from Georges Bank to Chesapeake Bay

(Figures 2 and 3; Bigelow and Schroeder 1953; Dery and

Livingstone 1982; Chang 1990). Windowpane generally

inhabit shallow waters (< 110 m) with sand to sand/silt or

mud substrates; they are most abundant from depths of 1-2

m (Warfel and Merriman 1944) to depths < 56 m (Thorpe

1991). They occur in most of the bays and estuaries south

of Cape Cod, including Chesapeake Bay (Hildebrand and

Schroeder 1928), Delaware Bay (de Sylva et al. 1962),

Sandy Hook Bay (Wilk and Silverman 1976), Raritan Bay

(Wilk et al. 1996), Long Island Sound (Moore 1947;

Gottschall et al., in review), and Narragansett Bay

(Jefferies and Johnson 1973). North of Cape Cod,

windowpane inhabit nearshore waters, but their

occurrence in estuaries is not well documented. Table 1

presents a qualitative summary of the distribution and

relative abundance of windowpane life history stages in

estuaries from Maine to Virginia (Jury et al. 1994; Stone

et al. 1994).

EGGS

The eggs are buoyant and spherical, with a diameter

of 0.9-1.4 mm), and a single oil globule 0.2-0.3 mm in

diameter (Wheatland 1956). At a typical spawning

temperature of 11o

C, hatching occurs in eight days (Miller

et al. 1991).

LARVAE

At hatching, windowpane larvae are approximately 2

mm long (Fahay 1983; Able and Fahay 1998). Flexion

begins at about 5.5 mm TL (Fahay 1983); eye

transformation during metamorphosis begins at about 6.5

mm TL (Colton and Marak 1969; Fahay 1983). The body

is darkly pigmented over most of its length. As

development proceeds, the body becomes deeper and

more laterally compressed. Fin ray formation is complete

at about 11.5 mm TL. Details of larval development are

provided by Moore (1947).

JUVENILES

The body is oval and wider (60-70% SL) than in

other left-eyed flounders. The body and fins are heavily

pigmented in larger young-of-the-year; smaller individuals

are characterized by broad alternating dark and light

bands. The mouth is large, extending to the eye or beyond

and the lateral line is arched over the pectoral fin (Figure

1; Able and Fahay 1998). The growth patterns of young

juveniles in estuaries and on the shelf vary with the timing

of spawning. Fish spawned in the spring grow quickly

and reach sizes of 11-19 cm TL by September, about four

months after spawning. By the following spring, most fish

of this cohort are larger than 16 cm TL. Fish spawned in

the autumn are 4-7 cm TL in December and reach 18-21

cm TL by the following October (Morse and Able 1995;

Able and Fahay 1998).

ADULTS

Windowpane attain a maximum total length of about

46 cm (Scott and Scott 1988). Few age and growth

studies of windowpane have been conducted (Moore

1947; Shelton 1979; Thorpe, 1991). It is a fast growing

species and spring and summer is the period of greatest

growth (Moore 1947).

REPRODUCTION

Gonadal development indices (Wilk et al. 1990) and

egg and larval distributions (Colton and St. Onge 1974;

Smith et al. 1975; Colton et al. 1979; Morse et al. 1987)

indicate that spawning occurs throughout most of the year.

Spawning begins in February or March in inner shelf

waters, peaks in the Middle Atlantic Bight in May, and

extends onto Georges Bank during the summer (Able and

Fahay 1998). Spawning also occurs in the southern

portion of the Middle Atlantic Bight in the autumn (Smith

et al. 1975). There is a split spawning season in the

central Middle Atlantic Bight with peaks in the spring and

autumn (Morse and Able 1995; Able and Fahay 1998).

Evidence for a split spawning season is available for

Virginia and North Carolina (Smith et al. 1975), for Long

Island Sound, New York (Wheatland 1956), and for Great

South Bay, New York (Dugay et al. 1989; Monteleone

1992). Gonad development indicated that split spawning

off New Jersey and New York peaks in May and in

September (Wilk et al. 1990). However, neither
Page 2

Perlmutter (1939) nor Smith et al. (1975) found evidence

for a split spawning season in Long Island Sound or in

oceanic waters north of Virginia. Colton and St. Onge

(1974) collected larvae on Georges Bank from July to

November but found no indication of a split spawning

season.

Some spawning may occur in the high salinity

portions of estuaries in the Middle Atlantic Bight,

including Great South Bay, New York (Monteleone

1992), Sandy Hook Bay, New Jersey (Croker 1965),

inside Hereford Inlet, New Jersey (Allen et al. 1978), and

in the coastal habitats of the Carolinas (Wenner and

Sedberry 1989). Windowpane spawn in the evening or at

night (Ferraro 1980) on or near bottom at temperatures

ranging from 6-21o

C (Bigelow and Schroeder 1953;

Wheatland 1956; Smith et al. 1975). Most spawning

(70%) was found at bottom water temperatures between

8.5-13.5o

C; spawning stopped off Virginia and North

Carolina when water temperatures exceeded 15o

C (Smith

et al. 1975).

Sexual maturity occurs at 3-4 years of age when about

50% of females that are 22 cm TL are sexually mature.

Females grow larger and faster than males after sexual

maturity (O’Brien et al. 1993).

FOOD HABITS

Juvenile and adult windowpane feed exclusively on

mysid shrimps in Johns Bay, Maine (Hacunda 1981).

Stomach content data collected during Northeast Fisheries

Science Center (NEFSC) bottom trawl surveys indicate

windowpane feed on small crustaceans (e.g., mysids and

decapod shrimp) and various fish larvae including hakes

and tomcod, as well as their own species (Langton and

Bowman 1981; Figure 4).

PREDATION

Spiny dogfish, thorny skate, goosefish, Atlantic cod,

black sea bass, weakfish and summer flounder are major

predator of windowpane, primarily juveniles.

MIGRATION

Juveniles that settle in shallow inshore waters move

to deeper offshore waters as they grow (Klein-MacPhee,

in prep.). Juveniles and adults may migrate to nearshore

or estuarine habitats in the southern Middle Atlantic Bight

in the autumn (Figures 2 and 3), however, juveniles are

probably not adequately sampled by standard Northeast

Fisheries Science Center trawl gear (Morse and Able

1995). Juveniles inhabiting Georges Bank (< 60 m)

undergo seasonal movements to deeper waters along the

southern flank of the Bank occur during late autumn, as

bottom temperatures drop, and overwintering occurs in

deeper areas until late spring (Figure 2).

STOCK STRUCTURE

Fish stocks are generally defined as having a fixed

spawning ground, a definite spawning season, and a

consistent migratory or movement pattern. Nonetheless,

spawning in windowpane occurs throughout most of the

year (April-December) and is closely linked to bottom

temperature (Colton and St. Onge 1974; Smith et al.

1975; Colton et al. 1979; Morse et al. 1987). Thus, stock

structure of windowpane could not clearly be identified.

However, the species is managed as two stocks: a northern

stock, Gulf of Maine-Georges Bank region, and a southern

stock, southern New England-Middle Atlantic Bight

region.

HABITAT CHARACTERISTICS

The habitat characteristics and preferences of

windowpane are summarized in Table 2. The methods

used to collect the fishery-independent survey data used in

this characterization are summarized in Reid et al. (1999).

EGGS

Windowpane eggs were collected at integrated water

column temperatures of 5-20o

C. Most eggs were

collected at 4-16o

C in spring (March-May), 10-16oC in

summer (June-August) and 14-20o

C in autumn

(September-November) in depths < 70 m (Figure 5).

LARVAE

Larvae settle to the bottom at approximately 10 mm

TL (Bigelow and Schroeder 1953). However, individuals

collected on Georges Bank may be planktonic up to 20

mm (Morse and Able 1995). Based on collections from

southern New Jersey, it appears that settlement of springspawned individuals occurs in estuaries and on the shelf,

while settlement of autumn-spawned individuals occurs

primarily on the shelf. Larvae are found throughout the

polyhaline portion of estuaries in the spring, but primarily

on the shelf in the autumn (Morse and Able 1995).

The maximum abundance of small larvae (< 5 mm

TL) occurred from 15-19o

C in areas south of Georges

Bank and at 14-15o

C on Georges Bank. Windowpane

larvae were collected during the NEFSC Marine

Resources Monitoring, Assessment and Prediction

(MARMAP) ichthyoplankton survey at integrated water

column temperatures of 5-20o

C, but mostly at 3-14oC in

spring, 10-17o

C in summer, and 13-19oC in autumn in
Page 3

water < 70 m deep (Figure 6).

JUVENILES

Juveniles were collected on the continental shelf

throughout the year during NEFSC bottom trawl surveys

(Figure 2) at a wide range of bottom temperatures (3-

25o

C) and depths (5-125 m). Juveniles were most

abundant at bottom temperatures of 4-7o

C in spring and

14-16o

C in autumn at depths < 50 m (Figure 7).

Juveniles inhabiting Massachusetts inshore waters

(Figure 8) were most abundant at 5-12o

C in spring and 12-

19o

C in autumn, and at depths < 20 m (Figure 9).

Windowpane were common in the Rhode Island

bottom trawl survey in Narragansett Bay; juveniles were

caught throughout the bay in all seasons with no

indication of seasonal differences. Juveniles were

captured at most bottom depths but showed a preference

for depths < 30 m in warmer bottom water temperature

periods (9-25o

C), and depths > 30 m in colder water

temperatures (1-8o

C). They occurred at a wide range of

bottom water temperatures: winter (1-8o

C), spring (1-

15o

C), summer (13-25oC), and autumn (10-21oC).

The bottom trawl survey in the Hudson-Raritan

estuary showed that juveniles were fairly evenly

distributed throughout the estuary, but they were most

abundant in the deeper channels in winter and summer

(Figure 10; Wilk et al. 1996). For all seasons combined,

juveniles were collected at bottom temperatures of 0-

24o

C, at depths < 25 m with salinities of 15-33 ppt, and

dissolved oxygen (DO) levels of 2-13 mg/l (Figure 11;

Wilk et al. 1996). Juvenile windowpane were most

abundant at bottom water temperatures of 5-23o

C, at

depths of 7-17 m, at salinities of 22-30 ppt, and DO levels

of 7-11 mg/l (Figure 11; Wilk et al. 1996).

ADULTS

The windowpane is a year-round resident off southern

New Jersey and probably in the Gulf of Maine (KleinMacPhee, in prep.). Adult windowpane tolerate a wide

range of temperatures (0-26.8o

C) and temperature may

control the northern extent of the species as well as its

local abundance (Moore 1947). In the Northeast Fisheries

Science Center bottom trawl survey (Figure 3), adults

were caught at bottom temperatures of 4-8o

C and depths <

75 m in spring and at 12-18o

C and depths < 50 m in

autumn (Figure 7).

Data from the Massachusetts inshore trawl survey

(Figure 8) indicated that most adults were caught south of

Cape Cod during spring at bottom temperatures of 9-13o

C

and at depths < 15 m. In autumn, adults were more

widely distributed and were caught at bottom temperatures

of 9-19o

C and depths < 30 m (Figure 9).

Adults were caught throughout Narragansett Bay in

all seasons with no apparent seasonal shift in abundance.

Adults preferred deeper waters (> 30 m) in cold bottom

water temperature periods (1-8o

C) and remained in a

shallow water (< 30 m) in the warmer bottom water

temperature periods (9-23o

C).

The bottom trawl survey in Long Island Sound found

that juvenile and adult windowpane were most abundant

in spring (April-June) (Figure 12; Gottschall et al., in

review). In spring, they were caught at bottom

temperatures of 3-18o

C, at salinities of 21-31 ppt, and at

depths < 60 m. The distribution pattern in autumn

(September-November) was similar to the pattern in

spring, but abundance was reduced (Figure 12). In

autumn, windowpane adults were caught at bottom

temperatures of 8-23o

C, at salinities of 18-32 ppt, and at

depths < 50 m (Gottschall et al., in review).

Adults were fairly evenly distributed throughout the

Hudson-Raritan estuary, but they were more abundant in

deeper channels in the summer (Figure 10; Wilk et al.

1996). For all seasons combined, adults were collected at

bottom temperatures of 0-24o

C, at depths < 25 m, at

salinities of 15-33 ppt, and DO levels of 2-13 mg/l (Figure

11; Wilk et al. 1996).

Adult windowpane occur primarily on sand substrates

off southern New England and the Middle Atlantic Bight,

but are frequently caught on mud grounds in the Gulf of

Maine (Langton et al. 1994). Adults are euryhaline; they

occur at salinities of 5.5-36.0 ppt (Tagatz 1967).

Windowpane are sensitive to hypoxic conditions; few

were collected where DO concentrations were < 3 mg/l,

presumably because they avoid such conditions (Howell

and Simpson 1994).

Adult windowpane may travel along the coast for

considerable distances; in one case, they moved 129 km in

three months (Moore 1947). These movements may play

an important role in the intermingling of local populations

(Klein-MacPhee, in prep.).

In a species association study using NEFSC

groundfish survey bottom trawl data, windowpane

commonly occurred with yellowtail flounder (Limanda

ferruginea), ocean pout (Macrozoarces americanus), and

little skate (Raja erinacea) during spring (Colvocoresses

and Musick 1984). In autumn, windowpane were more

widely distributed across the shelf and occurred with

yellowtail flounder, little skate, northern searobin

(Prionotus carolinus), and spiny dogfish (Squalus

acanthias).

GEOGRAPHICAL DISTRIBUTION

The windowpane is distributed from the Gulf of St.

Lawrence to Cape Hatteras, North Carolina, but it is most

common south of Nova Scotia (Figure 13). The largest

catches occur on Georges Bank.
Page 4

EGGS

Windowpane eggs have been collected in several

studies (Colton and St. Onge 1974; Smith et al. 1975;

Colton et al. 1979; Morse et al. 1987; Berrien and

Sibunka 1999). Windowpane egg distributions from

NEFSC MARMAP ichthyoplankton surveys are

summarized in Figure 14. Eggs were collected at 16% of

the stations sampled; primarily at depths < 40 m between

Georges Bank and Cape Hatteras. Eggs densities were

generally low in the Gulf of Maine. Eggs were collected

in nearshore shelf waters in the Middle Atlantic Bight

from February to November. Egg densities peaked in

May and October. Eggs were present on Georges Bank

from April through October and density peaked during

July-August.

LARVAE

The spatial distribution of windowpane larvae

collected in NEFSC MARMAP ichthyoplankton surveys

is summarized in Figure 15. More than 99% of the larvae

collected were 2-10 mm TL. Peak densities of recentlyspawned larvae (2-4 mm TL) occurred in the southern

Middle Atlantic Bight in May and November, and on

Georges Bank in July-October (Morse and Able 1995;

Figure 15). The larval distribution mirrors that of the eggs

in space and time.

JUVENILES

The spatial pattern of abundance for juvenile

windowpane on the continental shelf in the Middle

Atlantic Bight is similar to the spatial pattern for larvae

(Morse and Able 1995). Juveniles occur nearshore in the

Middle Atlantic Bight (< 40 m) and off southern New

England (< 50 m) throughout the year (Figure 2). On

Georges Bank, the spatial distribution of densities of

juveniles differs between spring and autumn (Wigley and

Gabriel 1991), and adults migration is similar to juveniles.

Spatial distribution of juveniles in the Gulf of Maine

shows low densities in nearshore areas in spring and

autumn.

In the Hudson-Raritan estuary, juveniles were fairly

evenly distributed throughout the estuary, but juveniles

were most abundant in the deeper channels in winter and

summer (Figure 10; Wilk et al. 1996).

ADULTS

The spatial distribution of adults on the continental

shelf (Figure 3) is similar to the distribution of juveniles

(Figure 2). Adults may migrate to nearshore or estuarine

habitats in the southern Middle Atlantic Bight during

spring through autumn. Adults on Georges Bank also

show seasonal movements to deeper waters from late

autumn through spring similar to juveniles. Adults in the

Gulf of Maine use nearshore waters during the spring and

autumn. The spring aggregation of adult windowpane in

Nantucket Sound and on Nantucket Shoals is evident in

the Massachusetts trawl survey (Figure 8). This

aggregation suggests spawning or feeding activities;

however, there is no supporting information on the

densities of eggs, larvae, or prey organisms.

STATUS OF THE STOCKS

The NEFSC autumn bottom trawl survey has been

used to estimate the relative abundance and biomass of

windowpane (Hendrickson 1998). The abundance index

for the Gulf of Maine-Georges Bank region generally

increased from the mid-1960s to a peak in 1984 and then

declined (Figure 16). The abundance index for the

southern New England-Middle Atlantic Bight region

declined sharply from 1963 to 1975 and has remained

relatively low since then (Figure 16).

The windowpane is managed by the New England

Fishery Management Council under the Multispecies

Fishery Management Plan (NEFMC 1993). This plan

defines overfishing for windowpane when the 3-year

moving average of the autumn stock abundance index falls

below the lowest quartile of the time series. Accordingly,

windowpane stock in the Gulf of Maine-Georges Bank is

considered to be fully exploited (Hendrickson 1998) while

southern New England-Middle Atlantic Bight stock is

overfished (National Marine Fisheries Service 1997;

Hendrickson 1998).

The distributions of windowpane were compared

between a period of high abundance (1984-1988) and a

period of low abundance (1992-1996) based on the

autumn Northeast Fisheries Science Center bottom trawl

survey (Figure 17). The spatial extent of adults and

juveniles was similar between the two periods.

RESEARCH NEEDS

• Studies to determine if the windowpane population is

a unit stock or multiple stocks (e.g., genetics, otolith,

cohort analysis).

• Windowpane spawning times and locations, and

spawning habitat requirements (e.g., high salinity).

• Studies (tagging, more efficient gear to catch younger

fish) to determine seasonal use of estuaries (residency

during colder months) and nearshore waters.

• Habitat requirements for windowpane eggs, larvae,

and juveniles.

• Growth rate studies.
Page 5

ACKNOWLEDGMENTS

This review was prepared with assistance from

members of the Essential Fish Habitat team. Jeffrey Cross,

Luca Cargnelli, Michael Fahay, Sara Griesbach, Frank

Steimle, Anne Studholme and Joseph Vitaliano provided

valuable suggestions, reference materials, survey maps,

histograms, food habit figures, and figures for fishery

resources. Judy Berrien, Rande Ramsey-Cross, and Claire

Steimle searched the reference literature. Special thanks to

Lisa Hendrickson and others at Woods Hole Laboratory,

and Michael Pentony and others at New England Fishery

Management Council, for providing critical reviews of an

earlier version of this report.

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Page 7

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Page 8

Table 1. Summary of the distribution and abundance of windowpane in North Atlantic and Mid-Atlantic estuaries based

on Jury et al. (1994) and Stone et al. (1994).

Estuary Adults Spawning

Adults

Juveniles Larvae Eggs

TMS TMS TMS TMS TMS

 Passamaquoddy Bay cc cc cc cc cc

 Englishman/Machias Bays c c c c c c c c c c

 Narragaugus Bay c c c c c c c c c c

 Blue Hill Bay c c c c c c c c c c

 Penobscot Bay ca ca ca ca ca

 Muscongus Bay c c c c c c cc cc

 Damariscotta Bay cc cc cc cc cc

 Sheepscot Bay c c cc c c cc cc

 Kennebec/Androscoggin Rivers c c c c c c cc cc

 Casco Bay c c c c c c c c c c

 Saco Bay c c c c c c c c c c

 Wells Harbor nz c c nz c c nz c c nz c c nz c c

 Great Bay rc rc rc rc rc

 Merrimack River r nz r nz r nz r nz r nz

 Massachusetts Bay nz nz c nz nz c nz nz c nz nz c nz nz c

 Boston Harbor nz c c nz c c nz c c nz c c nz c c

 Cape Cod Bay nz a a nz c c nz a a nz c c nz c c

 Waquoit Bay nz c c nz c c nz c c nz c c nz c c

 Buzzards Bay nz a a nz c c nz a a nz c c nz c c

 Narragansett Bay r a a c c r a a c c c c

 Long Island Sound r h h h h rhh cc hh

 Connecticut River h nz h nz h nz c nz h nz

 Gardiners Bay nz c c nz c c nz c c nz c c nz c c

 Great South Bay nz a a nz a a nz a a nz c c nz a a

 Hudson River/Raritan Bay r c c r c r c c r c c r r c

 Barnegat Bay h h hh h h hh hh

 New Jersey Inland Bays h h hh h h hh hh

 Delaware Bay a a a a

 Delaware Inland Bays nz c c nz c c nz c c nz c c nz c c

 Chincoteague Bay nz nz c nz nz nz nz c nz nz nz nz

 Chesapeake Bay mainstream cc cc r

 Chester River nz nz nz nz nz

 Choptank River nz nz nz nz nz

 Patuxent River nz nz nz nz nz

 Potomac River nz nz nz nz nz

 Tangier/Pocomoke Sounds nz c nz nz nz nz c nz nz nz nz nz

 Rappahannock River nz nz nz nz nz

 York River nz nz r nz nz nz

 James River r nz nz r nz nz nz

Relative Abundance

h = highly abundant, a = abundant,

c = common, r = rare, blank = not

present, n = no data presented,

* = no data available, nz = particular

zone not present

Data Reliability for Life Stages

Highly Certain = Bold and Underlined Text

Moderately Certain = Bold Text

Reasonable Inference = Normal Text

Tidal Zones

T = Tidal Fresh 0.0-0.5 ppt

M= Mixing Zone 0.5-25 ppt

S = Seawater Zone > 25 ppt
Page 9

Table 2. Summary of life history and habitat parameters for windowpane.

Life

Stage

Size and

Growth

Geographic

Location

Time of Year Habitat Substrate Temperature,

Salinity and DO

Prey/Predators

Eggs 1 0.9-1.4 mm Middle Atlantic

Bight

Georges Bank

Feb-July

Sept-Nov

May-Oct

Planktonic; less than

70 m

Not applicable Water column temp:

6-14o

C spring

10-16o

C summer

14-20o

C autumn

Eaten by adults of

own and other

species.

Larvae 2 2-10 mm Middle Atlantic

Bight

Georges Bank

Feb-July

Sept-Nov

May-Oct

Planktonic; less than

70 m

Not applicable Water column temp:

3-14o

C spring

10-17o

C summer

13-19o

C autumn

Prey on copepods

and other

zooplankton.

Eaten by adults of

own and other

species.

Juveniles 3 < 22 cm TL Gulf of Maine

Georges Bank

Middle Atlantic

Bight

June-Oct

June-Oct

May-July

Oct-Nov

Nearshore bays and

estuaries; less than

50 m

Less than 50 m

(summer/autumn);

less than 75 m

(winter/spring)

Nearshore bays and

estuaries; less than

75 m

Muddy sediment in

the Gulf of Maine

Fine sandy

sediment in

Georges Bank

Fine sandy

sediment in New

England & Middle

Atlantic Bight

Bottom temp:

Offshore:

4-7o

C in spring;

14-16o

C in autumn

Inshore off MA:

5-12o

C in spring

1 2-19o

C in autumn

Hudson-Raritan Bay

0-24o

C

(15-33 ppt - Salinity)

(2-13 mg/l - DO)

Prey on polychaetes

and small

crustaceans,

especially mysids.

Eaten by adults of

own and other

species (spiny

dogfish, thorny

skate, goosefish,

cod).

Adults 4 ≥ 22 cm TL Gulf of Maine

Georges Bank

Middle Atlantic

Bight

Year-round Nearshore bays and

estuaries; less than

75 m

Less than 50 m

(summer and

autumn);

less than 75 m

(winter and spring)

Nearshore bays and

estuaries; less than

75 m

Muddy sediment in

the Gulf of Maine

Fine sandy

sediment on

Georges Bank

Fine sandy

sediment

in New England

and Middle

Atlantic Bight

Bottom temp:

Offshore:

4-8o

C in spring

12-18o

C in autumn

Inshore off MA

9-13o

C in spring

9-19o

C in autumn

Hudson-Raritan Bay

0-24o

C

(15-33 ppt - Salinity)

(2-13 mg/l - DO)

Prey on polychaetes,

small crustaceans

(mysids, decapod

shrimp) various

small fishes (hakes,

tomcod).

Eaten by adults of

various fishes (spiny

dogfish, thorny

skate, goosefish,

cod).

1

 Colton and St. Onge (1974), Smith et al. (1975), Colton et al. (1979), Morse et al. (1987), Berrien and Sibunka (1999) 2

 Moore (1947), Colton and Marak (1969), Morse and Able (1995)

3

 Moore (1947), Thorpe (1991), Morse and Able (1995), Wilk et al. (1996), Able and Fahay (1998), Klein-MacPhee (in prep.) 4

 Colvocoresses and Musick (1984), Morse and Able (1995), Wilk et al. (1996), Able and Fahay (1998), Gottschall et al. (in review), Klein-MacPhee (in prep.)
Page 10

Figure 1. The windowpane, Scophthalmus aquosus (from Bigelow and Schroeder 1953).
Page 11

Figure 2. Distribution and relative abundance of juvenile windowpane (< 22 cm) from NEFSC bottom trawl surveys,

spring (1968-1997), summer (1963-1995), autumn (1963-1996), and winter (1964-1997). 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].

Number/Tow

 1 to 10

 10 to 25

 25 to 50

 50 to 75

 75 to 100

NMFS Trawl Surveys

Spring 1968 - 97

Juveniles (<22cm)

Windowpane

NMFS Trawl Surveys

Summer 1963 - 95

Juveniles (<22cm)

= Absent

 = Present

Windowpane

Number/Tow

 1 to 25

 25 to 50

 50 to 100

 100 to 500

 500 to 1079

NMFS Trawl Surveys

Autumn 1963 - 96

Juveniles (<22cm)

Windowpane

NMFS Trawl Surveys

Winter 1964 - 97

Juveniles (<22cm)

 = Absent

 = Present

Windowpane
Page 12

Figure 3. Distribution and relative abundance of adult windowpane (≥ 22 cm) from NEFSC bottom trawl surveys, spring

(1968-1997), summer (1963-1995), autumn (1963-1996), and winter (1964-1997). 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].

Number/Tow

 1 to 25

 25 to 50

 50 to 100

 100 to 500

 500 to 1121

NMFS Trawl Surveys

Spring 1968 - 97

Adults (>=22cm)

Windowpane

NMFS Trawl Surveys

Summer 1963 - 95

Adults (>=22cm)

= Absent

 = Present

Windowpane

Number/Tow

 1 to 25

 25 to 50

 50 to 100

 100 to 500

 500 to 1133

NMFS Trawl Surveys

Autumn 1963 - 96

Adults (>=22cm)

Windowpane

NMFS Trawl Surveys

Winter 1964 - 97

Adults (>=22cm)

 = Absent

 = Present

Windowpane
Page 13

Figure 4. Abundance (percent occurrence) of the major types of prey identified in the stomachs of juvenile and adult

windowpane collected during NEFSC bottom trawl surveys during 1973-1980 and 1981-1990. Note that the use of 20

cm as the segregation size between juvenile and adults differs from the actual size generally used (22 cm); this is an

artifact of the diet database that summarizes results in 10 cm length intervals. 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].

Arthropoda 85.5%

Fish 6.6%

Platyhelminthes 2.6%

Annelida 2.0%

All Other Prey 3.3%

Arthropoda 65.2%

Fish 17.5%

Unknown Animal Remains 5.1%

Miscellaneous 3.0%

Platyhelminthes 2.7%

Annelida 2.3%

All Other Prey 4.3%

Annelida 0.9%

Arthropoda 78.0%

Chaetognatha 4.6%

Fish 6.4%

Unknown Animal Remains 10.1%

Arthropoda 66.6%

Fish 23.0%

Annelida 0.8%

Unknown Animal Remains 6.5%

Chaetognatha 1.3%

All Other Prey 1.8%

1-20 cm

n = 134

20-40 cm

n = 535

1-20 cm

n = 107

21-50 cm

n = 816

a) 1973-1980

b) 1981-1990
Page 14

Figure 5. Percentage of windowpane eggs in relation to water column temperature (0-200 m, o

C) and bottom depth (m)

from NEFSC MARMAP ichthyoplankton surveys, February to November, 1978-1987 (all years combined). Open bars

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

standardized catches (number/10 m2

).

.

April

0

10

20

30

40

50

May

0

10

20

June

Percent

0

10

20

50

60

July

0

10

20

August

0

10

20

October

0

10

20

September

0

10

20

February

0

10

20

80

100

Stations

Egg Catch

March

0

10

20

30

40

November

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

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

0

10

20

30

40

April

0

10

20

60

80

October

0

10

20

30

40

50

60

September

0

10

20

30

50

60

May

0

10

20

30

60

70

June

Percent

0

20

80

100

July

0

10

20

30

40

August

0

10

20

30

40

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

50

March

0

10

20

30

80

100

February

0

10

20

90

100

Stations

Egg Catch
Page 15

Figure 6. Percentage of windowpane larvae in relation to water column temperature (0-200 m, o

C) and bottom depth (m)

from NEFSC MARMAP ichthyoplankton surveys, January to December, 1977-1987 (all years combined). Open bars

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

standardized catches (number/10 m2

).

January

0

10

20

30

Stations

Larva Catch

April

0

10

20

30

40

50

May

0

10

20

30

June

Percent

0

10

20

30

July

0

10

20

30

40

August

0

10

20

November

0

10

20

30

40

September

0

10

20

30

March

0

10

20

30

40

50

October

0

10

20

February

0

10

20

60

80

December

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

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

0

10

20

30

January

0

10

20

30

60

70

Stations

Larva Catch

April

0

10

20

60

80

December

Bottom Depth (m), Interval Midpoint

10

30

50

70

90

110

130

150

170

190

210

230

250

270

290

325

375

450

750

1250

1750

>2000

0

10

20

30

40

50

September

0

10

20

30

40

50

May

0

20

40

60

June

Percent

0

10

20

60

80

July

0

20

40

60

August

0

10

20

30

40

50

October

0

10

20

30

40

50

March

0

10

20

30

40

50

February

0

10

20

7080

November

0

20

40

60
Page 16

Figure 7. Percentage of juvenile and adult windowpane in relation to bottom water temperature and depth, based on

spring (1968-1997) and autumn (1963-1996) NEFSC bottom trawl surveys (all years combined). Open bars represent

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

standardized catches (number/10 m2

).

1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 0

10

20

30

40

1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 0

5

10

15

20

25

0

10

20

30

0

10

20

30

40

50

0

10

20

30

40

0

10

20

30

1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 0

4

8

12

16

20

1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 0

5

10

15

20

Juveniles 25 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)

NMFS Bottom Trawl Surveys

Windowpane
Page 17

Figure 8. The distribution and relative abundance of juvenile and adult windowpane from Massachusetts inshore trawl

surveys, spring and autumn 1978-1996 [see Reid et al. (1999) for details].

Windowpane

Mass. Inshore Trawl Survey

 Spring 1978 - 1996

 Juveniles (<22cm)

Number/Tow

 1 to 10

 10 to 25

 25 to 50

 50 to 100

 100 to 246

Windowpane

Mass. Inshore Trawl Survey

 Autumn 1978 - 1996

 Juveniles (<22cm)

Number/Tow

 1 to 10

 10 to 25

 25 to 50

 50 to 100

 100 to 133

Windowpane

Mass. Inshore Trawl Survey

 Spring 1978 - 1996

 Adults (>=22cm)

Number/Tow

 1 to 50

 50 to 100

 100 to 500

 500 to 1000

 1000 to 1661

Windowpane

Mass. Inshore Trawl Survey

 Autumn 1978 - 1996

 Adults (>=22cm)

Number/Tow

 1 to 10

 10 to 25

 25 to 50

 50 to 100

 100 to 133
Page 18

Figure 9. Percentage of juvenile and adult windowpane in relation to bottom water temperature and depth from the

spring and autumn Massachusetts inshore bottom trawl surveys (spring and autumn 1978-1996) for all years combined.

Open bars represent the proportion of all stations which were surveyed, while solid bars represent the proportion of the

sum of all standardized catches (number/10 m2

).

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

0

5

10

15

20

25

0

10

20

30

40

50

0

10

20

30

40

50

0

10

20

30

40

50

1 3 5 7 9 11 13 15 17 19 21 23 0

3

6

9

12

15

1 3 5 7 9 11 13 15 17 19 21 23 0

5

10

15

20

25

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

Windowpane
Page 19

Figure 10. Distribution and relative abundance of juvenile (< 22 cm) and adult (> 21 cm) windowpane collected during

spring, summer, autumn and winter in the Hudson-Raritan estuary from January 1992 to June 1997 [see Reid et al.

(1999) for details].

Staten

Island

NEW

YORK

NEW

JERSEY

No/Tow

 1 - 4

5 - 9

10 - 24

25 - 49

50 -105

Windowpane

Hudson-Raritan Estuary

Spring 1992 - 1997

Juveniles (<22 cm)

Staten

Island

NEW

YORK

NEW

JERSEY

No/Tow

 1 - 4

5 - 9

10 - 24

25 - 49

50 -105

Windowpane

Hudson-Raritan Estuary

Summer 1992 - 1996

Juveniles (<22 cm)

Staten

Island

NEW

JERSEY

No/Tow

 1 - 4

5 - 9

10 - 24

25 - 49

50 -105

Windowpane

Hudson-Raritan Estuary

Fall 1992 - 1996

Juveniles (<22 cm)

Staten

Island

NEW

YORK

NEW

JERSEY

No/Tow

 1 - 4

5 - 9

10 - 24

25 - 49

50 -105

Windowpane

Hudson-Raritan Estuary

Winter 1992 - 1997

Juveniles (<22 cm)
Page 20

Figure 10. cont’d.

Staten

Island

NEW

YORK

NEW

JERSEY

No/Tow

 1 - 4

5 - 9

10 - 24

25 - 49

50 -105

Windowpane

Hudson-Raritan Estuary

Spring 1992 - 1997

Adults (>21 cm)

Staten

Island

NEW

YORK

NEW

JERSEY

No/Tow

 1 - 4

5 - 9

10 - 24

25 - 49

50 -105

Windowpane

Hudson-Raritan Estuary

Summer 1992 - 1996

Adults (>21 cm)

Staten

Island

NEW

YORK

NEW

JERSEY

No/Tow

 1 - 4

5 - 9

10 - 24

25 - 49

50 -105

Windowpane

Hudson-Raritan Estuary

Fall 1992 - 1996

Adults (>21 cm)

Staten

Island

NEW

YORK

NEW

JERSEY

Windowpane

Hudson-Raritan Estuary

Winter 1992 - 1997

Adults (>21 cm)

No/Tow

 1 - 4

5 - 9

10 - 24

25 - 49

50 -105
Page 21

Figure 11. Percent frequency of juvenile and adult windowpane in relation to bottom water temperature, depth,

dissolved oxygen, and salinity in the Hudson-Raritan estuary, January 1992 to June 1997 (all years combined).

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

15 17 19 21 23 25 27 29 31 33 35

0

2

4

6

8

10

12

14

Juveniles (<22 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

40

15 17 19 21 23 25 27 29 31 33 35

0

5

10

15

20

Adults (>21 cm)
Page 22

Figure 12. Abundance and length frequency distributions of windowpane in Long Island Sound during spring and

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

Number/Tow

 1 to <50

 50 to <100

 100 to <250

 250 to <500

 500 to <628

Number/Tow

 1 to <25

 25 to <50

 50 to <100

 100 to <200

 200 to <424

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

Windowpane

Windowpane
Page 23

Figure 13. Distribution and abundance of windowpane from Newfoundland to Cape Hatteras during 1975-1994. 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 24

Figure 14. The distribution and abundance of windowpane eggs collected from February to November, 1978-1987

during NEFSC MARMAP ichthyoplankton surveys [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

Windowpane

Eggs

MARMAP Ichthyoplankton Surveys

61-cm Bongo Net; 0.505-mm mesh

February to November; 1978 to 1987

Number of Tows = 8476; with eggs = 1341

Eggs / 10m2

1 to <10

10 to <100

100 to <1000

1000 to <3890

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

35

36

37

38

39

40

41

42

43

44

45

Windowpane

Eggs

MARMAP Ichthyoplankton Surveys

61-cm Bongo Net; 0.505-mm mesh

February; 1978 to 1987

Number of Tows = 459; with eggs = 1

None

1 to 4

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

Windowpane

Eggs

MARMAP Ichthyoplankton Surveys

61-cm Bongo Net; 0.505-mm mesh

March; 1978 to 1987

Number of Tows = 853; with eggs = 59

None

1 to <10

10 to <100

100 to 137

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

Windowpane

Eggs

MARMAP Ichthyoplankton Surveys

61-cm Bongo Net; 0.505-mm mesh

April; 1978 to 1987

Number of Tows = 1020; with eggs = 88

None

1 to <10

10 to <100

100 to <1000

Eggs / 10m2

1000 to 1777
Page 25

Figure 14. 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

Windowpane

Eggs

MARMAP Ichthyoplankton Surveys

61-cm Bongo Net; 0.505-mm mesh

May; 1978 to 1987

Number of Tows = 1085; with eggs = 231

None

1 to <10

10 to <100

100 to <1000

Eggs / 10m2

1000 to 1728

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

35

36

37

38

39

40

41

42

43

44

45

Windowpane

Eggs

MARMAP Ichthyoplankton Surveys

61-cm Bongo Net; 0.505-mm mesh

June; 1978 to 1987

Number of Tows = 709; with eggs = 125

None

1 to <10

10 to <100

100 to <1000

Eggs / 10m2

1000 to 3890

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

35

36

37

38

39

40

41

42

43

44

45

Windowpane

Eggs

MARMAP Ichthyoplankton Surveys

61-cm Bongo Net; 0.505-mm mesh

July; 1978 to 1987

Number of Tows = 781; with eggs = 173

None

1 to <10

10 to <100

100 to 739

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

Windowpane

Eggs

MARMAP Ichthyoplankton Surveys

61-cm Bongo Net; 0.505-mm mesh

August; 1978 to 1987

Number of Tows = 863; with eggs = 158

None

1 to <10

10 to <100

100 to 544

Eggs / 10m2
Page 26

Figure 14. 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

Windowpane

Eggs

MARMAP Ichthyoplankton Surveys

61-cm Bongo Net; 0.505-mm mesh

September; 1978 to 1987

Number of Tows = 747; with eggs = 146

None

1 to <10

10 to <100

100 to 883

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

Windowpane

Eggs

MARMAP Ichthyoplankton Surveys

61-cm Bongo Net; 0.505-mm mesh

October; 1978 to 1987

Number of Tows = 1044; with eggs = 221

None

1 to <10

10 to <100

100 to <1000

Eggs / 10m2

1000 to 1029

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

35

36

37

38

39

40

41

42

43

44

45

Windowpane

Eggs

MARMAP Ichthyoplankton Surveys

61-cm Bongo Net; 0.505-mm mesh

November; 1978 to 1987

Number of Tows = 915; with eggs = 139

None

1 to <10

10 to <100

100 to 650

Eggs / 10m2
Page 27

Figure 15. The distribution and abundance of windowpane larvae collected from January to December, 1977-1987

during NEFSC MARMAP ichthyoplankton surveys [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 - December (1977 to 1987)

Windowpane Flounder

Number of Tows = 11438, with larvae = 1086

1 to < 10

10 to < 100

100 to 747

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 = 6

Number of Larvae / 10m2

None

1 to 5

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 = 3

Number of Larvae / 10m2

None

1 to < 10

10 to 15

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 6
Page 28

Figure 15. 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 = 5

Number of Larvae / 10m2

None

1 to 9

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 = 147

Number of Larvae / 10m2

None

1 to < 10

10 to 76

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 = 74

Number of Larvae / 10m2

None

1 to < 10

10 to 70

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 = 159

Number of Larvae / 10m2

None

1 to < 10

10 to < 100

100 to 747
Page 29

Figure 15. 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 = 139

Number of Larvae / 10m2

None

1 to < 10

10 to < 100

100 to 645

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 = 94

Number of Larvae / 10m2

None

1 to < 10

10 to < 100

100 to 199

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 = 264

Number of Larvae / 10m2

None

1 to < 10

10 to < 100

100 to 472

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 = 168

Number of Larvae / 10m2

None

1 to < 10

10 to < 100

100 to 607
Page 30

Figure 15. 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

December (1977 to 1987)

Number of Tows = 603, with larvae = 24

Number of Larvae / 10m2

None

1 to < 10

10 to 47
Page 31

Figure 16. Commercial landings (mt), bottom trawl survey indices (stratified mean catch per tow), and smoothed survey

indices (3 year moving average of first order autoregression model to compensate for inter-year variability) for

windowpane in the Gulf of Maine-Georges Bank region and the southern New England-Middle Atlantic Bight region.

Gulf of Maine - Georges Bank

1960 1965 1970 1975 1980 1985 1990 1995 2000

Landings (mt x 1000)

0

1

2

3

Stratified mean catch/tow (kg)

0

1

2

3

Commercial landings (mt)

Survey index (kg)

Smoothed survey index (kg)

Southern New England - Middle Atlantic

Year

1960 1965 1970 1975 1980 1985 1990 1995 2000

Landings (mt x 1000)

0.0

0.5

1.0

1.5

2.0

2.5

Stratified mean catch/tow (kg)

0.0

0.5

1.0

1.5

2.0

2.5

Commercial landings (mt)

Survey index (kg)

Smoothed survey index (kg)
Page 32

Figure 17. Distribution and abundance of juvenile (< 22 cm) and adult (≥ 22 cm) windowpane during a period of

relatively low abundance (1992-1996) and a period of relatively high abundance (1984-1988) from autumn NEFSC

bottom trawl surveys.

Number/Tow

 1 to <25

 25 to <50

 50 to <100

 100 to <200

 200 to <233

NMFS Trawl Surveys

Autumn 1992 - 96

Adults (>=22cm)

Low Abundance

Windowpane

Number/Tow

 1 to <25

 25 to <50

 50 to <100

 100 to <200

 200 to <233

NMFS Trawl Surveys

Autumn 1984 - 88

Adults (>=22cm)

High Abundance

Windowpane

Number/Tow

 1 to <25

 25 to <50

 50 to <100

 100 to <200

 200 to <267

NMFS Trawl Surveys

Autumn 1992 - 96

Juveniles (<22cm)

Low Abundance

Windowpane

Number/Tow

 1 to <25

 25 to <50

 50 to <100

 100 to <200

 200 to <267

NMFS Trawl Surveys

Autumn 1984 - 88

Juveniles (<22cm)

High Abundance

Windowpane
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

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all issues, the series employs thorough internal scientific review,

but not necessarily external scientific review. For most issues,

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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

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