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NOAA Technical Memorandum NMFS-NE-137: Essential Fish Habitat Source Document: Windowpane, Scophthalmus Aquosus, Life History and Habitat Characteristics
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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., I 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.

1I1.

Review and Evaluation of the 1994 Experimental Fishery in Closed Area 11 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., I1 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., I 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-I12345.

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., I table. NTIS Access. No. PB99-16501 1.

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.

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NOAA Technical Memorandum NMFS-NE-137 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.

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 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

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/ejh. 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 updateswill 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 Region's policy on the use of species names in all technical communications is generally to follow the American Fisheries Society's lists of scientific and common names for fishes (i. e., Robinsetal. 199 1 3), mollusks (i.e.,

Turgeon et al. 19981), and decapod crustaceans (i.e., Williams et al. 19891), and to follow the Society for Marine Mammalogy's guidance on scientific and common names for marine mammals (i.e., Rice 19 9 8d). 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 1998').

'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.

hTurgeon, 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. PubI. 26; 526 p.

cWilliams, A.B. (chair); Abele, L.G.; Felder, D.L.; Hobbs, H.H., Jr.; Manning, R.B.; McLaughlin, P.A.; Pdrez Farfante, I. 1989. Common and scientific names of aquatic invertebrates from the United States and Canada: decapod crustaceans. Amer. Fish. Soc. Spec. PubI. 17; 77 p.

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

'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 fishery-JAMES 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 Mid-Atlantic 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 NMFS-NE series.

JEFFREY N. CROSS, CHIEF ECOSYSTEMS PROCESSES DIvISION NORTHEAST FISHERIES SCIENCE CENTER

Page v Contents

Introduction...................

I Life History I

Habitat Characteristics 2

G eo g rap h ica l D istrib u tio n.........................................................................................................................................................................

3 Status of the Stocks 4

Research Needs 4

A c k n o w le d g m e n ts............................................................................................................

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...................................

II 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 ei 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 I 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 I I T, 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 6-2 1 C (Bigelow and Schroeder 1953; Wheatland 1956; Smith et al. 1975).

Most spawning (70%) was found at bottom water temperatures between 8.5-13.5"C; spawning stopped off Virginia and North Carolina when water temperatures exceeded 15'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 5-20TC.

Most eggs were collected at 4-16'C in spring (March-May), 10-16'C in summer (June-August) and 14-20°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 spring-spawned 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 15-19TC in areas south of Georges Bank and at 14-15'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 5-20'C, but mostly at 3-14'C in spring, 10-17TC in summer, and 13-19TC 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 (3-25TC) and depths (5-125 m).

Juveniles were most abundant at bottom temperatures of 4-7"C in spring and 14-16'C in autumn at depths < 50 m (Figure 7).

Juveniles inhabiting Massachusetts inshore waters (Figure 8) were most abundant at 5-12'C in spring and 12-19TC 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 (9-25"C), and depths > 30 m in colder water temperatures (1-8"C).

They occurred at a wide range of bottom water temperatures: winter (1-8"C),

spring (I-15°C), summer (13-25"C), and autumn (10-2 1°C).

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 0-24'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 5-23'C, at depths of 7-17 m, at salinities of 22-30 ppt, and DO levels of 7-11 mg/I (Figure 11; Wilk et al. 1996).

ADULTS The windowpane is a year-roundresident off southern New Jersey and probably in the Gulf of Maine (Klein-MacPhee, in prep.). Adult windowpane tolerate a wide range of temperatures (0-26.8'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 4-8'C and depths <

75 m in spring and at 12-18TC 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 9-13TC and at depths < 15 m.

In autumn, adults were more widely distributed and were caught at bottom temperatures of 9-19'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 (1-8°C) and remained in a shallow water (< 30 m) in the warmer bottom water temperature periods (9-23'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 3-18"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.8-230 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 0-24°C, at depths < 25 m, at salinities of 15-33 ppt, and DO levels of 2-13 mg/l (Figure 11; Wilk etal. 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 recently-spawned 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.

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.

Allen, D.M., J.P. Clymer, III, and S.S. Herman. 1978.

Fishes of the Hereford Inlet Estuary, southern New Jersey. Lehigh University, Dep. Biology, Center for Marine and Environmental Studies and The Wetlands Institute. 138 p.

Berrien, P. and J. Sibunka. 1999. Distribution patterns of fish eggs in the United States northeast continental shelf ecosystem, 1977-1987. NOAA Tech. Rep.

NMFS 145.310 p.

Bigelow, H.B. and W.C. Schroeder. 1953. Fishes of the Gulf of Maine. U.S. Fish Wild]. Serv. Fish. Bull., 53.

577 p.

Chang, S.

1990.

Seasonal distribution patterns of commercial landings of 45 species off the northeastern United States during 1977-88. NOAA Tech. Mem. NMFS-F/NEC-78. 130 p.

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. Nat]. Mar. Fish. Serv. Northeast Fish. Cent. Woods Hole Lab. Ref. Doc. 69-9. 43 p.

Colton, J.B. and J.M. St. Onge. 1974. Distribution of fish eggs and larvae in continental shelf waters, Nova Scotia to Long Island. Serial Atlas of the Marine Environment, Folio 23.

American Geographical Society, NY.

Colton, J.B., W.G. Smith, A.W. Kendall, Jr., P.L. Berrien, and M.P. Fahay. 1979. Principal spawning areas and times of marine fishes, Cape Sable to Cape Hatteras.

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

Colvocoresses, J.A. and J.A. Musick.

1984. Species associations and community composition of Middle Atlantic Bight continental shelf demersal fishes. Fish.

Bull. (U.S.) 82: 295-313.

Croker, R.A. 1965. Planktonic fish eggs and larvae of Sandy Hook estuary. Chesapeake Sci. 6: 92-95.

Dery, L. and R. Livingstone, Jr. 1982. Windowpane. In M.D. Grosslein and T.R.

Azarovitz eds.

Fish distribution. p. 114-116. MESA New York Bight Atlas Monograph

15. N.Y. Sea Grant Institute, Albany, NY.

de Sylva, D.P., F.A. Kalber, Jr., and C.N. Shuster. 1962.

Fishes and ecological conditions in the shore zone of the Delaware River Estuary, with notes on other species collected in deeper water.

University Delaware Mar. Lab. Inf. Ser. Publ. 5. 164 p.

Duguay, L.E., D.M. Monteleone, and C.-E; Quaglietta.

1989. Abundance and distribution of zooplankton and ichthyoplankton in Great South Bay, New York during the brown tide outbreaks of 1985 and 1986. In E. M. Cosper, V. M. Bricelj, and E. J. Carpenter. eds..

Novel phytoplankton blooms: causes and impacts of recurrent brown tides and other unusual blooms. p.

599-623. Coastal and Estuarine Studies No. 35.

Springer-Verlag, Berlin.

Fahay, M.P. 1983. Guide to the early stages of marine fishes occurring in the western North Atlantic Ocean, Cape Hatteras to the southern Scotian Shelf. J.

Northwest Atl. Fish. Sci. 4: 423 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.

Gottschall, K., M.W. Johnson, and D.G. Simpson. In review. The distribution and size composition of finfish, American lobster and long-finned squid in Long Island Sound based on the Connecticut Fisheries Division bottom trawl survey, 1984-1994.

NOAA Tech. Rep.

Hacunda, J.S.

1981.

Trophic relationships among demersal fishes in a coastal area of the Gulf of Maine.

Fish. Bull. (U.S.) 79: 775-788.

Hendrickson, L. 1998. Windowpane. In S.H. Clark ed.

Status of the fishery resources off the northeastern United States for 1998. p. 85-87. NOAA Tech. Mem.

NMFS-NE-115.

Hildebrand, S.F. and W.C. Schroeder. 1928. Fishes of Chesapeake Bay. Bull. U.S. Bur. Fish. 43(1): 366 p.

Howell, P. and D. Simpson. 1994. Abundance of marine resources in relation to dissolved oxygen in Long Island Sound. Estuaries 17: 394-402.

Jeffries, H.P.

and W.C.

Johnson.

1973.

Seasonal distributions of bottom fishes in the Narragansett Bay area: seven-year variations in the abundance of winter flounder (Pseudopleuronectes americanus). J. Fish.

Res. Board Can. 31: 1057-1066.

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 Assessment Division. Silver Spring, MD. 221 p.

Klein-MacPhee, G.

In preparation.

Windowpane Scophthalmus aquosus (Mitchill 1814). In B.B.

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Page 6 Schroeder's fishes of the Gulf of Maine. Smithsonian Institution Press, Washington, DC.

Langton, R.W. and R.E. Bowman. 1981. Food of eight northwest Atlantic pleuronectiform fishes. NOAA NMFS SSRF-749. 16 p.

Langton, R.W., J.B. Pearce, and J.A. Gibson eds. 1994.

Selected living resources, habitat conditions, and human perturbations of the Gulf of Maine:

Environmental and ecological considerations for fishery management. NOAA Tech. Mem. NMFS-NE-106. 70 p.

Miller, J.M., J.S. Burke, and G.R. Fitzhugh. 1991. Early life history patterns of the Atlantic North American flatfish: Likely (and unlikely) factors controlling recruitment. Neth. J. Sea Res. 27: 261-275.

Monteleone, D.M. 1992. Seasonality and abundance of ichthyoplankton in Great South Bay, New York.

Estuaries 15: 230-238.

Moore, E. 1947. Studies on the marine resources of southern New England. VI. The sand flounder, Lophopsetta aquosa (Mitchill); a general study of the species with special emphasis on age determination by means of scales and otoliths. Bull. Bingham Oceanogr. Collect. 11(3): 1-79.

Morse, W.W. and K.W. Able. 1995. Distribution and life history of windowpane, Scophthalmus aquosus, off the northeastern United States. Fish. Bull. (U.S.) 93:

675-693.

Morse, W.W., M.P. Fahay, and W.G. Smith. 1987.

MARMAP surveys of the continental shelf from Cape Hatteras, North Carolina to Cape Sable, Nova Scotia (1977-1984).

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2.

Annual distribution patterns of fish larvae. NOAA Tech. Mem. NMFS-F/NEC-47.215 p.

National Marine Fisheries Service.

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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.

[NEFMC] New England Fishery Management Council.

1993.

Final amendment

5 to the Northeast Multispecies Fishery Management Plan incorporating the supplemental environmental impact statement.

Vol. 1. September 1993. NEFMC. [Saugus, MA.]

361 p.

O'Brien, L., J. Burnett, and R.K. Mayo. 1993. Maturation of nineteen species of finfish off the northeast coast of the U. S., 1985-1990. NOAA Tech. Rep. NMFS 113. 66 p.

Perlmutter, A. 1939. A biological survey of the salt waters of Long Island, 1938. An ecological survey of young fish and eggs identified from tow-net collections. 28t.

Ann. Rep. N.Y. State Cons. Dep., Suppl., Pt. II: 11-71.

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.

Scott, W.B. and M.G. Scott. 1988. Atlantic fishes of Canada. Can. Bull. Fish. Aquat. Sci. 219. 731 p.

Shelton, S.T. 1979. The age, growth and food habits of the windowpane flounder, Scophthalmus aquosus (Mitchill), in the lower Cape Fear River estuary and adjacent ocean. M.S. thesis, East Carolina University, Greenville, NC. 42 p.

Smith, W.G., J.D. Sibunka, and A. Wells. 1975. Seasonal distributions of larval flatfishes (Pleuronectiformes) on the continental shelf between Cape

Cod, Massachusetts and Cape Lookout, North Carolina, 1965-1966. NOAA Tech. Rep. NMFS SSRF-691. 68 p.

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

Nelson, S.H. Jury, and M.E. Monaco. 1994. Distri-bution and abundance of fishes and invertebrates in Mid-Atlantic estuaries.

ELMR Rep.

No.

12.

NOAA/NOS Strategic Environmental Assessment Division, Silver Spring, MD. 280 p.

Tagatz, M.E. 1967. Fishes of the St. Johns River, Florida.

Q. J. Fla. Acad. Sci. 30(1): 25-50.

Thorpe, E.A. 1991. Aspects of the biology of windowpane flounder Scophthalmus aquosus, in the northwest Atlantic Ocean. M.S. thesis, Univ. Massachusetts, Amherst, MA. 101 p.

Warfel, H.E. and D. Merriman. 1944. Studies on the marine resources of southern New England. I. An analysis of the fish population of the shore zone. Bull.

Bingham Oceanogr. Collect. 9(2): 1-91.

Wenner, C.A.

and G.R.

Sedberry.

1989.

Species composition, distribution and relative abundance of fishes in the coastal habitat off the southeastern United States. NOAA Tech. Rep. NMFS-79. 49 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.

Wigley, S.E. and W.L. Gabriel. 1991. Distribution of sexually immature components of 10 northwest Atlantic groundfish species based on Northeast Fisheries Center bottom trawl surveys 1969-1986.

NOAA Tech. Mem. NMFS-F/NEC-80. 17 p.

Wilk, S.J., E.M. MacHaffie, D.G. McMillan, A.L.

Pacheco, R.A. Pikanowski, and L.L. Stehlik. 1996.

Fish, megainvertebrates, and associated hydrographic observations collected in the Hudson-Raritan Estuary, January 1992-December 1993. U.S. Natl. Mar. Fish.

Serv., Northeast Fish. Sci. Cent. Ref. Doc. 96-14. 95 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. 1976. Summer benthic fish

Page 7 fauna of Sandy Hook Bay New Jersey. NOAA Tech.

Rep. NMFS SSRF-698. 16 p.

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 Juveniles Larvae Eggs Adults T

M S

T M

S T

M S

T M

S T

M S

Passamaquoddy Bay c

c c

c C

C Englishman/Machias Bays c

c c

c Narragaugus Bay c

c c

c Blue Hill Bay c

c c

c Penobscot Bay c

a c

a' c

a c

a Muscongus Bay c

c c

c Damariscotta Bay c

c c

C C

C Sheepscot Bay c

C C

c c

C C

Kennebec/Androscoggin Rivers c

c c

c Casco Bay c

c c

c Saco Bay c

c c

c Wells Harbor nz c

c nz c

c Great Bay r

c r

C Merrimack River r

nz r

nz Massachusetts Bay nz nz g

nz nz c

Boston Harbor 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 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

r h

h c

c h

h Connecticut River h

nz h

nz c

nz h

nz Gardiners Bay nz c

c nz c

c Great South Bay nz a

a nz a

a nz c

c nz a

a Hudson River/Raritan Bay r

c c

r c

c r

c c

r r

c Barnegat Bay h h h

h h

h New Jersey Inland Bays h

h h

h Delaware Bay a

a a

a Delaware Inland Bays nz c

c nz c

c Chincoteague Bay nz nz g

nz nz nz nz c

nz nz nz nz Chesapeake Bay mainstream c

c C

C r

Chester River nz nzz 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 n.z nz James River r

nz nz r

nz nz nnz 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 Size and Geographic Time of Year Habitat Substrate Temperature, Prey/Predators Stage Growth Location Salinity and DO Eggs 0.9-1.4 mm Middle Atlantic Feb-July Planktonic; less than Not applicable Water column temp:

Eaten by adults of Bight Sept-Nov 70 m 6-14'C spring own and other 10-16"C summer species.

14-20'C autumn Georges Bank May-Oct Larvae 2-10 mm Middle Atlantic Feb-July Planktonic; less than Not applicable Water column temp:

Prey on copepods Bight Sept-Nov 70 m 3-14'C spring and other 10-1 7'C summer zooplankton.

13-19'C autumn Eaten by adults of Georges Bank May-Oct own and other species.

Juveniles

< 22 cm TL Gulf of Maine June-Oct Nearshore bays and Muddy sediment in Bottom temp:

Prey on polychaetes estuaries: less than the Gulf of Maine Offshore:

and small 50 m 4-70C in spring; crustaceans, 14-16"C in autumn especially mysids.

Eaten by adults of Georges Bank June-Oct Less than 50 m Fine sandy Inshore off MA:

own and other (summer/autumn);

sediment in 5-12'C in spring species (spiny less than 75 m Georges Bank I 2-19'C in autumn dogfish, thorny (winter/spring) skate, goosefish, cod).

Middle Atlantic May-July Nearshore bays and Fine sandy Hudson-Raritan Bay Bight Oct-Nov estuaries; less than sediment in New 0-24"C 75 m England & Middle (15-33 ppt - Salinity)

Atlantic Bight (2-13 mg/1 - DO)

Adults

>22 cm TL Gulf of Maine.

Year-round Nearshore bays and Muddy sediment in Bottom temp:

Prey on polychaetes, estuaries: less than the Gulf of Maine Offshore:

small crustaceans 75 m 4-8'C in spring (mysids, decapod 12-18"C in autumn shrimp) various small fishes (hakes, Georges Bank Less than 50 m Fine sandy Inshore off MA tomcod).

(summer and sediment on 9-13'C in spring Eaten by adults of autumn);

Georges Bank 9-19°C in autumn various fishes (spiny less than 75 m dogfish, thorny (winter and spring) skate, goosefish, Hudson-Raritan Bay cod).

Middle Atlantic Nearshore bays and Fine sandy 0-24'C Bight estuaries; less than sediment (15-33 ppt - Salinity) 75 m in New England (2-13 mg/I - DO) and Middle Atlantic Bight Colton and St. Onge (1974). Smith et al. (1975). Colton ei alt (1979). Morse et al. (1987), Berrien and Sibunka (1999) 2 Moore (1947), Colton and Marak (1969). Morse and Able (1995)

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

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 Windowpane NMFS Trawl Surveys Spring 1968 - 97 Juveniles (<22cm)

Number/Tow I 1o 10 S10 o 25 25 lo 50 50 :o 75 75 i 100 Windowpane NMFS Trawl Surveys Autumn 1963 - 96 Juveniles (<22cm)

Number/Tow I

I t, 25 25 to 50 50 to I(")

I I15) to 50) 5(X) to 1079 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].

Page 12 Windowpane Windowpane NMFS Trawl Surveys NMFS Trawl Surveys Spring 1968 - 97 Summer 1963 - 95 Adults (>=22cm)

Adults (>="'m 0 Presen

/"

¢"**"-.L_

x Absent

..,..:.C

'r4 NumbefrTow

" I t 25 25 lo 50

50 to (00 0

100 to 5M0 5(X) to 1121 Windowpane Windowpane NMFS Trawl Surveys NMFS Trawl Surveys AA d tsn( >1 2 2c 'm96

.j

/

W inter 1964 - 97.

Adumlts(2m Adults (>=22cm)

Number/Tow j-

I 2 to 250" 4

50(1( t 1133 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 (1 964-1997). 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 13 a) 1973-1980 1-20 cm n= 134 20-40 cm n = 535 Artlhropoda 85.5%

b) 1981-1990 Artlhropoda 78.0%

Arthropoda 65.2%

Al Other Prey 43%

Anmelida 2.3%

Platyhelminthes 2.7%

h Miscellaneonu 3

.0%

IUnkmmrtwfinrrl Rerrroimo5.1%

AlN Other Prey 3.3%

Anelda 2.0%

Platyhelrninthes 2.6%

1-20 cm na 107 21-50 cm n = 816 Arthropoda 66.6%

Annelida 09%

UnkownArnirral Remains 10.1%

All Other Prey 1.8%

Chaetognatha 1.3%

U n k no wn A n i e a l R e m ai n s 6.5 %

Annelida 0.8%

Fish 23.0%

Fish 6.4%

Cheetognatha 4.6%

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].

Page 14 910 20 1

40 20 0

30 20 I10 0

20 10 0

lo 60 50 20)

I

_A1 bruary 1

st,,,,

IEgg Caich 1~

March

~

April

~

May I

June

'I-if 3'£ U

I1

'em E631111L.

0 6

-l

~t'P"P Y-~-~-~-

July August

&a

¶a__

9 100 200 February*

Son, 100 20 March 208J U

,I*

Jn 30 70 800 2

August I10 3'

U September 00 October 70 2o JJ~

November B0oM 0'of.

BotmDph mItrvlMdon 2(l ll)

(1 0 ýLftAqwjnn 20 September Id Ji

[

October 203 November 2

4 6

8 1

12 14 16 19 20 22 24 26 28 Water-Column Temperature (0-200m, C)

Figure 5. Percentage of windowpane eggs in relation to water column temperature (0-200 m, '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/l10 m 2).

Page 15 C

03 C-20

~~January Sttin 30 20 February 30 March 30 April 20 20 May 20 June 40o,

I July 20 iAugust Flo J Ji g.l 30-20 September

'0" O"9 211 October*

o - -

30 November 20 302),0 i

llt December 0

2 4

6 8

10 12 14 16 10 20 22 24 26 20 Water-Column Temperature (0-200m, C) 6 3

2 54 3

2 I

1 C

03 03 6

4 2

9 6

2 6

4 2

4 3

2 1

5 3

3 2

4 2

2 0

Mm January February 0-0 March 30-0 April 0

041.

May

° 1June 60 50 0Oto July 0

0 0

DAugust 0-I 0 j'D September 0

October 0

November 0

December 04 Bottm Depth (in), Interval Midpoint Figure 6. Percentage of windowpane larvae in relation to water column temperature (0-200 m, °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/l10 M 2 ).

Page 16 Windowpane nI Stations NMFS Bottom Trawl Surveys 0

Catches 40-Juveniles 25 Adults

30.

Spring 205 Spring 30--

20" 1In" I 3 5 7 9 11 131517.1921 23252729 3 5 7 9 11131517192123252729 Bottom Temperature (C)

Bottom Temperature (C) 25-20 20' 16-Autumn Autumn 15-5 4-50-I 3 5 7 9 11131517192123252729 1 3 5 7 9 11 131517 192123252729 Bottom Temperature (C)

Bottom Temperature (C) 301 30-Spring Spring 201 20 Bottom Depth (in)

Bottom Depth (m) 50-40) 4[)-

Autumn 30-Autumn 30 20 2n-In" 10-

)

()-i U0ý0F Bottom Depth (m)

Bottom Depth (m)

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 m ).

Page 17 Windov Mass. Inshore a

C

~~Spring 1978-19 Juveniles

.n Is lo 2 tX.;........

mb.

-4 i

".'.* :-i..:.:

i a.**.

- .q "

<22cm)

r/low o10 (1 Windov Mass, Inshore T1 "Spring 1978 Adults (>=

, I;Z.

---I

50 1

J

~50C001

.V 4

xf Y-,..... W.

o50 to 1~

500 0 ) o 1661 Windowpane Mass. Inshore Trawl Survey Autumn 1978-1996 Adults (>=22cm)

I.o25

25 to 501 100 to 133 V

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 at. (1999) for details].

Page 18 20' 16" 12' 9.

4.

Juveniles AILhkI Windowpane Aass. Inshore Trawl Surveys Adults 3

Stations]

N Catches Spring 25' Spring 20' 15' 10'MU L

u 1 3 5

7 9

I1 13 15 17 19 21 23 Bottom Temperature (C) 30-Autumn 20" 10" 1

3 5

7 9

11 13 15 17 19 21 23 Bottom Temperature (C) 25 20' Spring 15" IO" 10' 5"

Bottom Depth (m) 50-40-Autumn 301 211 P P

,P P

qqqq 1 3 5

7 9

II 13 15 17 19 21 23 Bottom Temperature (C) 15-Autumn 12' 9-6' 3

1 3

5 7

9 11 13 15 17 19 21 23 Bottom Temperature (C) 50' 40 30 20' 10' Spring L

~n n n n..

Bottom Depth (m) 50' 40 30' 20' IH' Autumn Bottom Depth (m)

SL4 5

.ott N

k9 Depth S(m)

Bottom Depth (in)

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/lO mi2 ).

Page 19 NEW Windowpane

/

r^

Hudsmn-Raritan Estuary Spring 1 9 92-1997 Juveniles (<22 cm)

" D 9

W ial n

e

4.

NEW

-0 10.24 JERSEY M09) lo Figure 10. Distribution and relative abundance of juvenile (< 22 cm) and adult (> 21 cm) windowpane collected during spring, summer, autumn andwinter in the Hudson-Raritan estuary from January 1992 to June 1997 [see Reid et al.

( 1999) for details].

Page 20 NEW

.noaeYORK Wnopn Windowpane Windowpane Hudson-Raritan Estuary Hudson-Raritan Estuary 1

um o 992 19996 Spring 9

Summer 1992 - 1996 Adults (>21 cm)

Adults (>21 cm)

Staten

.0Staiens

- Islund o

.I.s n

v t

0,24 0

0-24 NEW~~~

249NW02-9 I.NEW oe to--

2 JERSEY 3

0isJRE nm EWNEW WindowpaneK Windowpane Hudson-Raritan Estuary Hudson-Raritan Estuary Hud n-Ranitan Est uary 1 7, Fall 1992 - 1996 Winter 1992-1997 Adults (>21 cm)

Adults (>21 cm)

S.

Adlt (21m)'

°e O*.

Ko K"

Statoen

I 0

Stin Island

' t Islandt e

° 0P

4FFN7T, No/Tos

.SO 0/024" NEW

-24 NEW 25 _ 49 JERSEY 50-10.5JERSEY

.d n u Figure 10. cont'd.

Page 21 Juveniles (<22 cm) 16]

14'

-i 12 Ci 10 8

6 4

2 UCatches

'Jr (U

P.

25 20 15 10 5-F.hFI F1I

~1.

I ml ml IIt/

0 2

4 6

8 10 12 14 16 18 20 22 24 26 Temperature (C) 0 1

2 3

4 5

6-7 8

9 10 11 12 13 Dissolved Oxygen (mg/I) 14' 12' 10 8-6 4

2 10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85 Depth (ft) 0* 15

17.

19 21 23 25 27 29 31 33 35 Salinity (ppt)

Adults (>21 cm) 16" 14" 12" 10" 4

6 4-2"1 H[Lk El] Stations Catches 1)

C-40 35 30 25 20" 15" 10 5.

mVLbrimFF1 I

DI m m

0 24 6

8 10 12 14 16 18 20 22 24 26 Temperature (C) 0 1 2 3

4 5

6 7

8 9 10 11 12 13 Dissolved Oxygen (mg/l) 4)UI-.

4) 35' 30 25 20 15.

10' 5.

4)

U 4) 10 15 2025 30354045 5055 6065 7075 80 85 Depth (ft) 15 17 19 21 23 25 27 29 31 33 35 Salinity (ppt)

Figure I1. 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).

Page 22 Fisheries Division

* *i,"**,!*

Long Island Sound Trawl Survey SPRING 1992 - 1997

°

'_L I*00.

1830

.I0 1.

00 U

Ch I,

State of Connecticut DEP Fi sheries Divisio L n Isa dSound Trawl Survey ",I*,,*,'

AUTUMN 1992-1997 1 1.45o 2*

Wge2A4indowpane i

IM 9W U

CZ 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].

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-table 1.html).

Page 24

45.

2 A_

I Windowpane Windowpane Eggs 44-Eggs MARMAP Icbthyoplankton Surveys ~/MARMAP Icbtbyoplankton Surveys 64-cm Bongo Ne; 0.505-mm mesh 43 61-cm Bongo Net; 0.505-mm mesh February to November; 1978 to 987 February; 1978 9o 1987 Numwer of Tows 459; with eggs I 42-Numbe ro"T-ws 9476; witheggo 1341 42-39-"

/39 EgEs//

IGm 17-*

1

Ii<IO 37 "I

o M

10to <I 000 36*

10001) <3890 36-76 75 74 73 72 71 7I70 69 61 7-66 65 76 75 74 73 72 71 70 69 64 8

67

ý6 45 45 Windowpane Windowpane 44,Eggs 4.Eggs MARM AP lchthyoplankton Surveys Eggs

/*¢* *',

MARMAP Ichthyoplankton Surveys M.M,

]S 61-cm Bongo Net; 0.505-mm m.esh 61-cm Bongo Net; 0.505m eh, March; 197810o1987 A pril; 197810o1987 Numr=of Tow.,

853; with eggs :59 Numbe otTows 1020: with egg% 89 39-391 37-I gs/10 Eggts/ l10t Noneo None I0 (otS100 0 tOtool<0 1000to 137 0

1Ooto<1000 36-

1000to 1777 35 35 76 75 74 73 72 71 70 69 64 67 66 65 76 75 74 73 72 71 70 69 68 67 66 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].

Page 25 76 75 74 72 7n 70 45 44-43-42-41-Windowpane

,Eggs MARMAP IchthyoplankEon Surveys 61-cm Bongo Net: 0.505-mm mesh July: 1978 to 1987 NumberfT.....= 781: with eggs=,73%x I

40-39 38-37jý 36~

Eggs / 1m0 None 1 ts-10(

0 t101-100 0

1C00t,739 72 71 701 69 69 67 66 Figure 14. cont'd.

Page 26 45--.

-* t t

Windowpane 41ý tEggs MARMAP tchthyoplankton Surveys 4J 61-cm Bongo Net; 0.505-mm mesh September: 197(8 to 1987 41 Eggs/ 10m No.e I tionl0

tOIouM 1001,,88X3 75 74 73 72 S1 70 69 64 67 66 Windowpane Eggs 44-43-42~

MARMAP Ichthyoplankton Surveys 61-cm Bongo Net; 0.505-mm mesh November; 1978 to 1987 Number of Tows =(915: with eggs = 13

/7.1 Ees / 1m None 1 1-to10 100 t, 650 74 73 7 2 7t 71 69 6

67 66 Figure 14. cont'd.

Page 27 45

~

245-Windowpane Flounder

Nuamhe, of Tow, = 434. wih 1.rvae =6 44-MARMAP Ichthyoplankton Surveys 44-61-cm Bongo Net 0.505-mm mesh January - December (1977 to 1987)

Numhor,-fT-.,

11439. woith I--u (106 4

42-1 42-41-41-

40) 40-39-3 38-Nue o L 10m2 3

Number of Larvae/IOm' Ito < 10 1 to5 37-I to < 100 37-1 (X) to 747 36-

.36-76 75 74 73 72 71 701 69 68 67 66 65 76 75 74 73 72 71 70 69 68 67 66 65 45-t L-_

.ii 52_4i February (1977 to 197)

March (1977 to 1987)

Number of Tows= 666. with 1-rvao = 3

)

Number of Tos = 103 1. with b-vae = 3 44-44-43-43:4 A5 42-42-40:

40 VN.

39-

-9 39-f Number of Larvae/ lin N,

38-Number of LarvaeI Om'lo None (None 37 /

oc)

< 10 1 to6 37l010tol

-5 37-36-36-35_: _

f 35-76 75 74 73 72 71 70 69 66 67 66 65 76 75 74 73 72 71 70 69 668 6 7 6 6 65 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].

Page 28 75 74 73 72 71

70) 69 6*

67 66 45-1 June (1977 to 1997)

Numhr o"I'

=

T 9,

. 93, wilh 1-=

74 44-1 43-41-Number of Larvae 10mrn None I to < l0 10 to 70(

74 7'3 2

71 70 69 6

67 66 Figure 15. cont'd.

Page 29 45-44-4 3-1 42-41-401 37-36-j 35 I I I

76 75 74 73 72 71 70 69 69 67 66 4 5..

..1 October (1977 to 1987)

N ujmbeo

,fTow = 1147. wilh larva e=264 44.t 43-42-41-40":

.39-1 Number of Larvae / I 0mý None I 1 toO< 10 S Oto< 100

100 to 472 37-j 36-1, 35 7--- 7 4

7' 7

7

)

6 6

7 7'6 7'5 74 73 72 "7

1 70 )

69 68 6 *7 66 Figure 15. cont'd.

/I Page 30 45-1..

J1 1

U December (1977 to 1987)

Numbee ol Tows = 603. with lI-v-e = 24 44-43-I.-

Number of Larvae / lOim None S

I to < 10 S

10to 47 37-76 75 74 73 72 71 70 69 68 67 66 Figure 15. cont'd.

Page 31 Gulf of Maine - Georges Bank 3

Commercial landings (mt)

Survey index (kg)

Smoothed survey index (kg) 00 X

C cz

_J 2

1-

.3

,20)

-0 0

11 1

0 I1...

I0...

I I9....

1 0

960 1965 1970 1975 1980 1985 1990 1995 2000 Southern New England - Middle Atlantic 2.5 80 0

0 X

CD) 0)

cz

-j 2.0 1.5 1.0 0.5 Commercial landinc Survey index (kg)

- Smoothed survey in A

2.5 gs (mt) ndex (kg) 2.0 1.5 1.0 0.5 0.0 1980 1985 1990 1995 2000

,a 0

-C Co C-a)

E "D

0.0 *H-1960 1965 1970 1975 Year 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.

Page 32 Windowpane AWindowpane NMFS Trawl Surveys NMFS Trawl Surveys Autumn 1992 - 96 Autumn 1984 - 88 Adults (>=22cm)

Adults (>=22cm)

Low Abundance High Abundance WA S. -

Number#Tow Number/Tow

/

I to <25 I to <25

25 to <ý50

25 to <50 o

50 to <10)0 6

50 to <100 10)0 to <1200

100 to <200)

/*200 to <233 is 200 to <1233 Windowpane 1

Windowpane NMFS Trawl Surveys NMFS Trawl Surveys Autumn 1992-96 Autumn 1984-88

/4 Juveniles (<22cm)

Juveniles (<22cm)

Low Abundance fHigh Abundance Number/Tow

.Number/Tow I 1to <25 I to 25 t, <50 25 to <50

50 wt <100 50 to <100 S1 200 to <200 200 to <267 1

200 to <2067 Figure 17. Distribution and abundance ofjuvenile (< 22 cm) and adult (> 22 cm) windowpane during a period of relatively low abundance (I 992-1996) and a period of relatively high abundance (1984-1988) from autumn NEFSC bottom trawl surveys.

ML072070389

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