NOAA Technical Memorandum NMFS NE 149 Essential Fish Habitat Source Document:

Scup, Stenotomus chrysops,

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:

Scup, Stenotomus chrysops, Life History and Habitat Characteristics Frank W. Steimle, Christine A. Zetlin, Peter L. Berrien, Donna L. Johnson, and Sukwoo Chang 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 149 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 Regionís policy on the use of species names in all technical communications is generally to follow the American Fisheries Societyís 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).

aRobins, 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.

bTurgeon, 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.

cWilliams, A. B. ( chair); Abele, L. G.; Felder, D. L.; Hobbs, H. H., Jr.; Manning, R. B.; McLaughlin, P. A.; PÈrez 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.

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

eCooper, 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 NMFSNE series.

JEFFREY N. CROSS, CHIEF ECOSYSTEMS PROCESSES DIVISION NORTHEAST FISHERIES SCIENCE CENTER Page v Contents Introduction............................................................................................................................................................................................... 1 Life History............................................................................................................................................................................................... 1 Habitat Characteristics .............................................................................................................................................................................. 4 Geographical Distribution ......................................................................................................................................................................... 5 Status of the Stocks ................................................................................................................................................................................... 7 Research Needs ......................................................................................................................................................................................... 8 Acknowledgments..................................................................................................................................................................................... 8 References Cited ....................................................................................................................................................................................... 8 Tables Table 1. Summary of life history and habitat characteristics for scup, Stenotomus chrysops................................................................. 14 Figures Figure 1. The scup, Stenotomus chrysops ( from Goode 1884) ............................................................................................................. 16 Figure 2. Abundance of the major items in the diet of scup collected during NEFSC bottom trawl surveys ....................................... 17 Figure 3. Abundance of scup eggs relative to water temperature and bottom depth from NEFSC MARMAP surveys ....................... 18 Figure 4. Abundance of scup larvae relative to water temperature and bottom depth from NEFSC MARMAP surveys ..................... 19 Figure 5. Seasonal abundance of scup relative to bottom water temperature and depth based on NEFSC bottom trawl surveys......... 20 Figure 6. Abundance of scup relative to bottom temperature, depth, DO, and salinity based on Hudson Raritan surveys .................. 22 Figure 7. Abundance of scup relative to bottom temperature and depth based on Massachusetts inshore bottom trawl surveys ......... 23 Figure 8. Seasonal abundance of scup relative to bottom temperature and depth from Narragansett Bay trawl surveys...................... 24 Figure 9. The distribution of scup from Newfoundland to Cape Hatteras ............................................................................................ 26 Figure 10. Distribution and abundance of scup eggs collected during NEFSC MARMAP surveys ....................................................... 27 Figure 11. Distribution and abundance of scup larvae collected during NEFSC MARMAP surveys..................................................... 29 Figure 12. Distribution and abundance of juvenile and adult scup collected during NEFSC bottom trawl surveys ............................... 30 Figure 13. Distribution and abundance of juvenile and adult scup in Massachusetts coastal waters ...................................................... 32 Figure 14. Seasonal distribution and abundance of juvenile and adult scup collected in Narragansett Bay ........................................... 33 Figure 15. Size frequency distribution of scup collected in Narragansett Bay during Rhode Island bottom trawl surveys .................... 35 Figure 16. Distribution, abundance, and size frequency of scup in Long Island Sound, from the Connecticut bottom trawl surveys.... 36 Figure 17. Seasonal distribution and abundance of juvenile and adult scup in the Hudson Raritan estuary........................................... 37 Figure 18. Commercial landings and NEFSC trawl survey indices for scup in southern New England and Middle Atlantic Bight ...... 39 Page 1 INTRODUCTION Scup ( Stenotomus chrysops Linnaeus 1766) ( Figure 1), is a temperate species that occurs primarily from Massachusetts to South Carolina, although it has been reported as far north as the Bay of Fundy and Sable Island Bank, Canada ( Bigelow and Schroeder 1953; Fritz 1965; Scott and Scott 1988) and as far south as Florida ( Morse 1978; Manooch 1984). The ` southern porgy' ( S. aculeatus) is referred to in a number of South Atlantic Bight studies and reviews ( e. g., Morse 1978; Powles and Barans 1980; Sedberry and Van Dolah 1984), but is not considered a separate species by the American Fisheries Society ( Robins et al. 1991) leading to some taxonomic confusion ( T. Munroe, National Systematics Laboratory, Smithsonian Institution, Washington, DC, personal communication). For example, Miller and Richards ( 1980) list S. chrysops and S. aculeatus as reef dwellers in the South Atlantic Bight. Although there can be some mixing of the Middle and South Atlantic Bight scup populations off North Carolina, the Middle Atlantic Bight population is treated separately here, because only this population appears to make extensive seasonal migrations and few fish tagged off New England or New York have been caught south of Cape Hatteras ( Nesbit and Neville 1935; Finkelstein 1971). Scup in the Middle Atlantic Bight population are commonly found during the summer in larger estuaries and in coastal waters; during the winter, they occur along the outer continental shelf to about 200 m ( 656 ft) and occasionally deeper. Beebe and Tee Van ( 1933) reported that scup were introduced to Bermuda, but the status of that introduction is unknown and probably unsuccessful ( B. Collette, National Systematics Laboratory, Smithsonian Institution, Washington, DC, personal communication). Archeological evidence suggests scup have been common in southern New England waters for several thousand years and were used as food by native Americans ( Waters 1967). The scup population in the Middle Atlantic Bight spawns along the inner continental shelf off southern New England from May through August with a peak in June to July. Larvae occur in coastal waters during the warmer seasons, feed upon small zooplankton, and are prey to a variety of planktivores, including medusae, crustaceans and fish. Larvae settle to the seafloor in coastal and estuarine waters when they are about 25 mm total length ( TL), but this event is poorly documented. During the summer and early fall, juveniles and adults are common in most larger estuaries and coastal areas in open and structured habitats where they feed on a variety of small benthic invertebrates. Scup begin to mature at 2 years of age ( Finkelstein 1969b) at about 15.5 cm fork length ( FL) ( O'Brien et al. 1993). Most fish are mature at 3 years and at 21 cm FL ( Gabriel 1998). In the last century, scup ³ 45 cm FL were reported ( Baird 1873) living to about 20 years and weighing about 2 kg ( Bigelow and Schroeder 1953). Currently, the population in the Middle Atlantic Bight is composed primarily of fish £ 7 years and £ 33 cm FL ( Northeast Fisheries Science Center 1997). Since the 1930s, there has been a significant decline in the average size of scup; small scup have slightly different habitat and prey requirements than larger scup ( Smith and Norcross 1968).

LIFE HISTORY The life history of scup is typical of most demersal fishes, with pelagic eggs and larvae, and a gradual transition to the demersal adult stage. As a temperate species, scup is at the northern limits of its range in the northeastern United States and migrates south in the winter to warmer waters south of New Jersey.

EGGS Scup eggs are small, 0.8 1.0 mm in diameter, and buoyant ( Kuntz and Radcliffe 1918; Wheatland 1956). They require two to three days ( 40 75 hrs) to hatch depending on temperature ( Griswold and McKenney 1984). Little else is known of this ephemeral stage.

LARVAE The newly hatched larvae are about 2.0 mm TL, pelagic, and depend on their yolk for about three days until they are about 2.8 mm TL ( Bigelow and Schroeder 1953) when active feeding begins. After reaching 15 30 mm TL in early July, the larvae become demersal in shoal waters ( Lux and Nichy 1971; Johnson 1978; MAFMC 1996; Able and Fahay 1998). Griswold and McKenney ( 1984) considered the larvae as juveniles when they grow to about 18 19 mm TL. There is no information available on habitat use or requirements during this transition period.

JUVENILES Able and Fahay ( 1998) noted that the smallest, young of the year ( YOY) individuals appeared in estuaries in June. In southern New England, juvenile scup grew to 5 to 10 cm FL by November ( Bigelow and Schroeder 1953; Gottschall et al., in review). Returning juveniles in the spring were about 10 13 cm FL ( Michelman 1988; Able and Fahay 1998). Growth of YOY scup is considered relatively slow ( Able and Fahay 1998). Michelman ( 1988) estimated daily growth of juveniles to be 0.84% of its dry wt/ day using a length frequency method and 0.93% of its dry wt/ day using a bioenergetics method. The growth production rates were Page 2 between 0.15 and 0.40 g of its dry wt/ m2 with a growth efficiency of about 24%. Growth rates and curves for juvenile scup were reported in several studies, see MAFMC ( 1996).

ADULTS Adult scup are common residents in the Middle Atlantic Bight from spring to fall and are generally found in schools on a variety of habitats, from open sandy bottom to structured habitats such as mussel beds, reefs or rough bottom. Smaller sized adult scup are common in larger bays and estuaries but larger sizes tend to be in deeper waters. Schools are reported to be size structured ( Morse 1978). Scup mature at about 2 years of age and 50% of both sexes are reported to be mature when they achieve a length of 15.5 cm FL ( O'Brien et al. 1993). Examining growth of male and female scup from the New York Bight ( the continental shelf bounded by southern Long Island and the New Jersey coast), Wilk et al. ( 1978) found no significant difference in the length weight relationships between sexes within the 113 361 mm FL range. The relationship for a larger sample of unsexed fish, 27 380 mm FL, was log W = log ( 5.022) + 3.169 log FL, where W is weight in grams and fork length ( FL) is in mm; similar relationships have been reported in MAFMC ( 1996). Growth in length is curvilinear between 10 38 cm FL corresponding to ages of about 1 to 13 years; growth is relatively rapid at 10 15 cm FL and declines with increasing size ( Penttila et al. 1989). Scup are members of an offshore wintering guild of fishes whose movements, habitats, and food habits generally coincide ( Musick and Mercer 1977; Colvocoresses and Musick 1984; Austen et al. 1994; Brown et al. 1996). This guild includes summer flounder ( Paralichthys dentatus), black sea bass ( Centropristis striata), northern searobin ( Prionotus carolinus), and smooth dogfish ( Mustelus canis) ( Gabriel 1992; Shepherd and Terceiro 1994). Although biological interactions among guild members can occur, slight differences exist in their environmental tolerances and habitat preferences ( Neville and Talbot 1964).

REPRODUCTION The mean fecundity of scup, 17.5 23.0 cm FL, is about 7,000 ( ± 4,860 SD) eggs per female ( Gray 1990). Scup spawn once a year beginning in the spring during the inshore migration ( Kendall 1973) when water temperatures are > 10 ° C. In eastern Long Island bays ( New York) and Raritan Bay ( New York New Jersey), spawning occurs in May and June ( Breder 1922; Finkelstein 1969a). Along coastal Rhode Island, spawning peaks in June ( O'Brien et al. 1993) and extends to August at temperatures of about 24 ° C ( Herman 1963). In southern Massachusetts, spawning fish occur in shoal waters < 10 m deep until late June, when they move into deeper waters ( MAFMC 1996). Most spawning occurs in southern New England from Massachusetts Bay south to the New York Bight, including eastern Long Island Sound, Peconic and Gardiners Bays, and Raritan Bay ( Goode 1884; Kuntz and Radcliffe 1918; Breder 1922; Nichols and Breder 1927; Permutter 1939; Bigelow and Schroeder 1953; Wheatland 1956; Richards 1959; Finkelstein 1969a; Sisson 1974; Morse 1978; Clayton et al. 1978). Able and Fahay ( 1998) noted that there has been no reported evidence of spawning in Block Island Sound ( Rhode Island), Great South Bay ( New York), the Hudson River estuary, and Great Bay ( New Jersey). Although Breder ( 1922) reported ripe scup in the Hudson Raritan estuary, more recent studies do not report the collection of scup eggs or larvae ( Croker 1965; Berg and Levinton 1985). Esser's ( 1982) note on scup spawning in the estuary was not referenced and is probably based on Breder ( 1922). Spawning has not been reported south of New Jersey ( Morse 1982); e. g., off Chesapeake Bay ( Hildebrand and Schroeder 1928; Pearson 1932). However, Berrien and Sibunka ( 1999) found eggs in this area between 1978 and 1987, although they were not abundant or widespread. Although scup are common in the spring off Maryland and Virginia, Eklund and Targett ( 1990) did not observe spawning over hard bottom reef habitat. The scup they observed appeared to be migrants since few remained as summer residents in the study area. Ferraro ( 1980) suggested that scup spawn in the morning in Peconic Bay, Long Island, unlike most fish that generally spawn in the evening or at night. Scup usually spawn over weedy or sandy areas and fertilization is external with no parental care ( Morse 1978). Scup appear to refrain from feeding during spawning ( Baird 1873; Bigelow and Schroeder 1953; Morse 1978). Spawning can fail in some years, e. g., 1958 ( Edwards et al. 1962), even though, based on landings data, spawning stocks are near peak abundance ( MAFMC 1996). The relationship of this apparent spawning failure to environmental or habitat variables is unknown. Scup spawning coincides temporally with that of several other fish, including weakfish ( Cynoscion regalis), tautog ( Tautoga onitis), and northern searobin ( Morse 1978).

FOOD HABITS Although food habits data for scup larvae are not available, rearing experiments suggest that the larvae feed on small zooplankton ( Griswold and McKenney 1984). In Long Island Sound, juvenile scup feed during the day, principally on polychaetes ( e. g., maldanids, nephthids, nereids, and flabelligerids), epibenthic amphipods and other small crustaceans, mollusks, and Page 3 fish eggs and larvae ( Bowman et al. 1987). Copepods and mysids are important to post larvae and early juveniles, while bivalve mollusks are more commonly eaten by larger fish ( Richards 1963b; Bowman et al. 1987; Michelman 1988). Allen et al. ( 1978) reported amphipods, polychaetes, copepods, and other small crustaceans were eaten by a small sample of juvenile scup in southern New Jersey, which is consistent with Northeast Fisheries Science Center ( NEFSC) data [ Figure 2; see Reid et al. ( 1999) for a discussion of NEFSC food habitats data]. Michelman ( 1988) reported that scup only eat when they are in a school and the relative importance of major prey taxa varies seasonally. Baird ( 1873) reported prey were " rooted out of the sand or mud." Juvenile and adult scup near an artificial reef in lower Delaware Bay ate a mix of hard surface epifauna and sand bottom infaunal prey, including amphipods ( caprellids and others), razor clams ( Ensis directus), hydroids, blue mussels ( Mytilus edulis), anemones, and mysids ( F. Steimle, unpublished data). In Raritan Bay, scup 9 12 cm FL ate a variety of benthic infaunal and epifaunal invertebrates including polychaetes, copepods, small mollusks, and hydroids; dietary composition varied among areas within the bay ( Steimle et al., in review). Michelman ( 1988) estimated that juvenile scup in Narragansett Bay ( Rhode Island) consumed 0.6 1.7 g dry wt/ m2 of benthic prey between June 1 and September 30. The daily food ration of juvenile scup was 3.49 3.99% of dry body weight ( depending on method used), or about 5% of their body weight per day. Adult scup are also benthic feeders and forage on a variety of prey, including small crustaceans ( including zooplankton), polychaetes, mollusks, small squid, vegetable detritus, insect larvae, hydroids, sand dollars, and small fish ( Goode 1884; Nichols and Breder 1927; Hildebrand and Schroeder 1928; Bigelow and Schroeder 1953; Oviatt and Nixon 1973; Maurer and Bowman 1975; Morse 1978; Sedberry 1983; Figure 2). As scup grow, their diets include larger prey. Bowman et al. ( 1976) found that polychaetes were more important in the diets of scup off southern New England and anthozoans were more important in the Middle Atlantic Bight. Sedberry ( 1983) reported that during the fall migration off New Jersey scup fed mainly on amphipods, polychaetes, and to a lesser extent on decapod crustaceans, copepods, snails, and other small invertebrates. Adults also prey on small benthic invertebrates, although feeding and growth appear to be reduced during the winter. At times and in certain areas, scup diets overlap those of red hake ( Urophycis chuss) and, depending on scup size, those of silver hake ( Merluccius bilinearis) and Gulf Stream flounder ( Citharichthys arctifrons) ( Sedberry 1983). Langton ( 1982) found that although the diets of scup overlapped those of several other demersal species, there was little prey overlap with cod ( Gadus morhua) or silver hake off New England, even though they have similar benthic diets. Jeffries and Terceiro ( 1985) hypothesized that an expanding scup population in Narragansett Bay seemed to replace the winter flounder ( Pseudopleuronectes americanus) because both species have similar diets; if abundance of winter flounder were reduced, more prey could be available for benthic feeding species such as scup. This dietary similarity was also found in a recent fish food habit study in Hudson Raritan Bay ( Steimle et al., in review). During inshore residency, scup gradually accumulate food reserves from the spring into the fall. The mean caloric content increases from 24.2 kj/ g ash free dry weight of whole scup in the spring to 28.1 kj/ g ash free dry weight in the fall ( Steimle and Terranova 1985). This stored energy can support the extra demands of migration, reduced feeding in winter, and gonadal development. Feeding may be minimal during the winter because there is so little growth ( Bigelow and Schroeder 1953).

PREDATION AND MORTALITY Larvae are probably preyed on by a variety of planktivores, including medusae, crustaceans, and fishes. Small or juvenile scup are heavily preyed on by bluefish ( Pomatomus saltatrix), Atlantic halibut ( Hippoglossus hippoglossus), cod, various sharks, striped bass ( Morone saxitilus), weakfish, goosefish ( Lophius americanus), silver hake, and other coastal fish predators ( Baird 1873; Smith 1898; Jensen and Fritz 1960; Schaefer 1970; Morse 1978; Sedberry 1983). Baird ( 1873) reported that cod ate large numbers of small scup on Nantucket Shoals in late November. Wading and diving shorebirds are also potential predators during the summer. The NEFSC bottom trawl survey data on food habits lists the following species as predators of scup: dusky shark ( Carcharhinus obscurus), sandbar shark ( C. plumbeus), smooth dogfish, spiny dogfish ( Squalus acanthias), Atlantic sharpnose shark ( Rhizoprionodon terraenovae), Atlantic angel shark ( Squatina dumeril), Atlantic torpedo ( Torpedo nobiliana), bluntnose stingray ( Dasyatis say), silver hake, bluefish, summer flounder, black sea bass, weakfish, northern stargazer ( Astroscopus guttatus), goosefish, inshore lizardfish ( Synodus foetens), and king mackerel ( Scomberomorus cavalla). Another potential source of mortality is disease. Disease can be initiated by direct epidermal exposure or through feeding on contaminated prey. Scup had fin rot in the degraded inner New York Bight and Hudson Raritan estuary ( Mahoney et al. 1975). Benthic invertebrate prey commonly eaten in the New York Bight were contaminated with several toxic heavy metals ( Steimle et al. 1994).

MIGRATION As inshore water temperatures decline to < 8 9oC in Page 4 the winter, scup leave inshore waters and move to warmer waters on the outer continental shelf south of the Hudson Canyon off New Jersey and along the coast from south of Long Island to North Carolina in depths ranging from 75 185 m ( Morse 1978; Bowman et al. 1987). Juveniles follow adults to wintering areas on the mid to outer continental shelf south of Long Island, although some remain in larger and deeper estuaries during warmer winters. During migration, scup move south along the coast ( within the 18 m isobath) and offshore ( Hamer 1970) as coastal bottom water temperature declines below 10oC. Phoel ( 1985) reported that scup migrated south of Cape Hatteras to about Cape Fear ( North Carolina) in the winter and spring ( he assumed one species and no population mixing). With rising water temperatures in the spring, scup return inshore. Larger fish arrive first followed by schools of subadults, which have been reported to appear off southern New England slightly later ( Sisson 1974). The fish reach Chesapeake Bay by April ( Hildebrand and Schroeder 1928) and southern New England by early May ( Baird 1873; Perlmutter 1939; Neville and Talbot 1964; Finkelstein 1971). It has been suggested that the population moves in schools of similarly sized individuals during migration and perhaps at other times as well ( Baird 1873; Hildebrand and Schroeder 1928; Neville and Talbot 1964; Sisson 1974; Morse 1978). Fish that arrive inshore early can be caught in pockets of residual cold water and can become inactive or dormant ( Kessler 1966).

STOCK STRUCTURE Although the Middle Atlantic Bight population was once considered to be two stocks, i. e., southern New England and New Jersey ( Edwards et al. 1962; Neville and Talbot 1964; Hamer 1970; Morse 1978). More recent analysis found that the evidence for this segregation was weak. Pierce ( 1981) suggested that the apparent segregation of two stocks in the Middle Atlantic Bight could be an artifact of the temporary location of separate winter water masses containing temperatures acceptable to scup; in most years this water mass separation is lacking or less influential. Scup is presently considered a single stock in the Middle Atlantic Bight ( Pierce 1981; Mayo 1982).

HABITAT CHARACTERISTICS Scup are a temperate, demersal species that use several benthic habitats from open water to structured areas for feeding and possibly for shelter ( Table 1). Their distribution changes seasonally as fish migrate from estuaries to the edge of the continental shelf as water temperatures decline in the winter and return from the edge of the continental shelf to inshore areas as water temperatures rise in the spring. Some reports on scup habitat use and distribution may be biased by the type of collection gear used and the habitats in which they can be deployed effectively. For example, most surveys use towed nets that are appropriate for open bottom but not for rough, structured habitats that scup are known to use such as mussel beds, rock rubble, or reefs.

EGGS Scup eggs are commonly found in larger bodies of coastal waters such as bays and sounds in and near southern New England during spring and summer. Lebida ( 1969) reported eggs were relatively abundant in Buzzards Bay ( Massachusetts) from May through June at water temperatures of 8.5o to 23.7oC, which is similar to their distribution in Connecticut and Rhode Island estuaries ( Herman 1963). Eggs hatched in about 70 75 hrs at 18oC and 40 54 hrs at 21 22oC ( Griswold and McKenney 1984); they may not develop normally at temperatures below 10oC ( Bigelow and Schroeder 1953). Few scup eggs were collected in the NEFSC Marine Resources Monitoring, Assessment and Prediction ( MARMAP) ichthyoplankton survey [ see Reid et al. ( 1999) for survey methods]. The few survey tows that collected eggs were made during May August when integrated water column temperatures were between 11o and 23oC ( Figure 3). Their occurrence at 23oC probably represents eggs collected off Maryland Virginia during the summer. Most eggs were collected in generally < 50 m ( Figure 3).

LARVAE Larval scup are pelagic and occur in coastal waters during warmer months. Larvae were collected in the more saline parts of Long Island Sound and eastern Long Island bays, Narragansett Bay, Buzzards Bay, Vineyard Sound, and Cape Cod Bay from May through September at water temperatures of 14 22oC; the greatest densities occurred at 15 20oC ( Fish 1925; Wheatland 1956; Pearcy and Richards 1962; Herman 1963; Scherer 1984; MAFMC 1996). Herman ( 1963) found larvae when water temperatures were 20.0 23.5oC. The optimum for rearing larvae in the laboratory is 18oC ( Lawrence 1979). The NEFSC MARMAP larval data indicate a peak in abundance at 17oC at depths < 50 m ( Figure 4).

JUVENILES During warmer months, juvenile scup live inshore in a variety of coastal habitats and can dominate the overall fish population in most larger estuarine areas during that period. In Rhode Island, YOY scup have been collected Page 5 in intertidal and subtidal habitats, over sand, silty sand, shell, mud, mussel beds and eelgrass ( Zosteria marina) ( Baird 1873). Although Gottschall et al. ( in review) noted that 1 year old scup were found on various types of sediment during warmer months in Long Island Sound, Richards ( 1963a) reported collecting more juvenile scup in a sandy habitat 9 m deep than at a 17 m deep muddy area of the sound. Scup were also collected in the smaller coastal bays of Delaware ( Derickson and Price 1973). However, scup were not common in shoreline seine or throw trap surveys in vegetated and unvegetated habitats in Chesapeake Bay, Long Island Sound, or New Jersey estuaries ( Greeley 1939; Warfel and Merriman 1944; Briggs and O'Connor 1971; Himchak 1982; Weinstein and Brooks 1983; Sogard 1989; Sogard and Able 1991). While little is known about the specific habitats occupied in winter when juvenile scup reside offshore, their winter spring distributions indicate that they occur in habitats ranging from relatively flat, open, sandy silty bottoms to the head of submarine canyons, and other areas with topographical relief and varying sediments ( Wigley and Theroux 1981). The presence of structure can be important to scup. Gray ( 1990) and Auster et al. ( 1991, 1995) noted that juveniles use biogenic depressions in the sediments off southern New England in the fall; the size of the depression was directly related to the size of the fish. Juveniles can use biogenic depressions, sand wave troughs, and possibly mollusk shell fields for shelter in winter. Their poor growth during colder months ( Bigelow and Schroeder 1953) suggests inactivity and possibly an increased need for shelter. Juvenile scup have been collected at water temperatures ranging from 5 27oC [ Figures 5 8; see Reid et al. ( 1999) for survey methods]. This is slightly below the thermal maximum of 30.2 35.6oC ( depending on acclimation) reported by Everich and Gonzalez ( 1977). The modes of highest relative abundance shift from about 10oC in the spring to peaks at 16oC and 22oC from summer to fall, except in Narragansett Bay ( Figure 8) and Long Island Sound where the bimodality was unclear. In Long Island Sound, where juveniles dominate the population, they were collected at bottom temperatures of 7 18oC in the spring and 15 22oC in the fall at salinities of 25 31 ppt. Subadults, which usually follow the migrations of adults south during the fall, have been killed by sudden cold spells in shallow New England bays ( Baird 1873; Sherwood and Edwards 1902; Morse 1978). However, from 1971 to 1975, juveniles over wintered in Long Island Sound ( Thomson et al. 1978). In the Hudson Raritan estuary, juveniles were collected at temperatures ranging from 9o to 26oC, at salinities ranging from 18 to 33 ppt, and dissolved oxygen ( DO) levels > 4 mg/ l ( Figure 6). From summer through fall, YOY and age 1+ scup were found in many tidal bays, sounds, and coastal areas primarily north of Maryland at depths within the 38 m (< 125 ft) contour ( Morse 1978; Figures 6 8). In Raritan Bay, juvenile scup were most commonly collected at depths between about 5 and 12 m ( 15 to 35 ft) ( Figure 6).

ADULTS Adult habitats are similar to those used by juveniles, including soft, sandy bottoms, on or near structures, such as rocky ledges, wrecks, artificial reefs, and mussel beds in euryhaline areas ( Briggs 1975a; Eklund 1988; MAFMC 1996). In Long Island Sound, scup exhibit a strong preference for mixed sand and mud sediments ( Gottschall et al., in review), which are probably rich in small benthic prey ( Reid et al. 1979). Similar to juveniles, the specific habitats used by adult scup during the winter or during migration are not known. The areas in which they have been found can include a variety of habitat types that differ in sediment composition, availability of food, and structure or relief ( Wigley and Theroux 1981; Steimle 1990). Adult scup also occurred at bottom water temperatures of 6 27oC ( Figures 5 8). Their winter distribution appears to be mostly limited by the 7oC isotherm, their lower preferred limit ( Neville and Talbot 1964). Magnuson et al. ( 1981) reported that scup may aggregate north of transient Gulf Stream frontal boundaries off Cape Hatteras, at least in the fall when the temperature differential was about 8oC ( 25.6o vs. 17.1oC). However, there are taxonomic uncertainties about the species of Stenotomus involved. Although scup are considered a demersal species, they have been observed at the water surface ( Bigelow and Schroeder 1953). Off Massachusetts ( Figure 7) and in Narragansett Bay ( Figure 8), most adults were collected in spring through fall at depths < 30 m ( 100 ft). In New Jersey, they were reported to aggregate within the 20 m depth coastal zone as they began their offshore southerly movements ( MAFMC 1996). Adult scup in the Hudson Raritan estuary were collected at salinities ranging primarily from 20 to 31 ppt ( Figure 6), which is consistent with salinity associations in Long Island Sound ( Gottschall et al., in review). Similar to juveniles in the Hudson Raritan estuary, most adults were collected at DO levels ³ 4mg/ l ( Figure 6).

GEOGRAPHICAL DISTRIBUTION Scup is a temperate species and north of Cape Hatteras the population is restricted to water temperatures above 6oC ( Figure 9). Postlarval scup migrate to stay within acceptable thermal limits as bottom water temperatures in the northeast decline in winter. Page 6 EGGS Scup eggs have been collected primarily in coastal waters off southern New England where abundance can range up to 1000 eggs/ 10 m2 of sea surface ( Berrien and Sibunka 1999) but samples containing > 100 eggs/ 10 m2 were rare during the NEFSC MARMAP survey ( Figure 10) when stock abundance was relatively low ( MAFMC 1996). Eggs were collected primarily during June and July from inshore waters off southern New England; few eggs were collected on the continental shelf from May to August ( Berrien and Sibunka 1999). Patchy occurrences were recorded from mid shelf in the Chesapeake Bight from May through August ( Figure 10). Since the NEFSC MARMAP surveys did not sample waters < 10 m and excluded most coastal bays, it is probable that eggs are more abundant and widely distributed in nearshore areas. Wheatland ( 1956) reported that in eastern Long Island Sound and nearby bays, eggs were variably abundant from year to year from May to August with peaks in June and July. According to Stone et al. ( 1994), scup eggs were common or abundant in the saline parts of coastal bays from southern Cape Cod to Long Island Sound, eastern Long Island, and the Hudson Raritan estuary. In contrast, Merriman and Sclar ( 1952) did not find eggs in Block Island Sound, along the south shore of Long Island, or in coastal waters or bays to the south. Interestingly, Able and Fahay ( 1998) note that there has not been a verified collection of scup eggs within southern New England estuaries since Sisson ( 1974). North of Cape Cod, scup eggs have been recorded in southern Cape Cod Bay from June to August ( 1974 1976), possibly transported from Buzzards Bay through the Cape Cod Canal ( Scherer 1984). There have been other reports of eggs in Massachusetts Bay suggesting that spawning occurs there ( MAFMC 1996).

LARVAE Larval distribution is also limited and even more conjectural than for eggs. Although Kendall ( 1973) noted the offshore occurrence of larvae from Virginia to Cape Cod and in estuaries from Delaware Bay to Buzzards Bay, the NEFSC MARMAP surveys collected < 5 larvae/ tow, mostly inshore ( about 30 m) off Rhode Island in July ( Figure 11). However, larvae can be more abundant in shallow, nearshore waters since Stone et al. ( 1994) reported them in the same areas as eggs; i. e., from southern Cape Cod to Long Island Sound and in the Hudson Raritan estuary. Despite these reports, Able and Fahay ( 1998) noted that like the eggs there has been no verified collection of scup larvae in southern New England estuaries since Sisson ( 1974). Cowen et al. ( 1993) did not collect scup larvae in coastal or shelf waters of the New York Bight during July and August 1988, nor were they common in bays or estuaries south of Long Island ( Pearson 1932; Massman et al. 1961; de Sylva et al. 1962; Dovel 1967, 1981; Scotton 1970; Pacheco and Grant 1973; Himchak 1982; Morse 1982; Olney 1983; Berg and Levinton 1985; Monteleone 1992; Stone et al. 1994) or in the surf zone ( D. Clark, U. S. Army Corps Engineers, Vicksburg, MS, personal communication). This is surprising since some of these areas; e. g., Delaware Bay, are important juvenile nurseries ( de Sylva et al. 1962). Clayton et al. ( 1978) reported the occurrence of larvae in Rocky Point in northwestern Cape Cod Bay, which, as with eggs, could have been transported through the Cape Cod Canal from Buzzards Bay ( Scherer 1984). Based on the presence of eggs and larvae, there is a possibility that scup can spawn in Massachusetts Bay ( MAFMC 1996).

JUVENILES In contrast with the conflicting reports and uncertainty in the spatial extent and abundance of scup eggs and larvae, juveniles have been collected inshore and offshore from New England to the Chesapeake Bay area. In fact, the saline areas of Narragansett Bay, Long Island Sound, Raritan Bay, and Delaware Bay are important nursery areas ( Richards 1963a; Abbe 1967; Oviatt and Nixon 1973; Werme et al. 1983; Michelman 1988; Gray 1990; MAFMC 1996; Wilk et al. 1997; Gottschall et al., in review). Reports of the coastal occurrence of juvenile scup date back to the last century. Smith ( 1894) reported that they were abundant from Hyannis, Massachusetts to Barnegat, New Jersey in 1891 and Moore ( 1894) indicated they were common only as far south as New Jersey. More recent reports indicate that during warmer months, juvenile scup were common from the intertidal zone to about 30 m in more saline (> 15 ppt) portions of bays and estuaries and along the inner continental shelf of the Middle Atlantic Bight from about May to November ( Smith 1898; Breder 1922; Kendall 1973; Werme et al. 1983; Bowman et al. 1987; Szedlmayer and Able 1996; Gottschall et al., in review). The changes in seasonal distribution are reflected in the results of the NEFSC bottom trawl surveys in which juveniles occurred offshore in winter and spring, inshore in summer, and were concentrated in near coastal waters through fall ( Figure 12). Young of the year fish are locally abundant north of Cape Cod ( Clayton et al. 1978), especially in the fall ( Lux and Kelly 1982). However, this is not reflected in the Massachusetts trawl survey that indicated higher concentrations south of the Cape in spring and fall ( Figure 13). Juveniles were common in Narragansett Bay ( Figure 14) and Long Island Sound ( Figure 16) in summer and fall. Zawacki and Briggs ( 1976) routinely seined juveniles on the north shore of Page 7 Long Island from July through October. Gottschall et al. ( in review) reported that YOY scup ( approximately 4 cm FL) were first collected in Long Island Sound in August and became numerically dominant in the catch by September; 1 year old juveniles were collected in April. However, other surveys of Long Island estuaries or surf zones did not support these findings ( Schaefer 1967; Briggs 1975b). The occurrence of juveniles in coastal bays and estuaries south of Long Island is temporally and spatially variable. In Raritan Bay, juveniles were abundant in spring and summer; a few were collected in the fall and were not collected in winter ( Figure 17). While juveniles occur in the larger bays; e. g., Raritan and Delaware Bays ( de Sylva et al. 1962; Werme et al. 1983), they seldom occur in smaller coastal lagoons such as Barnegat Bay ( New Jersey), tributaries of the Hudson Raritan estuary, or the ocean surf zone ( Marcellus 1972; Howells and Brundage III 1977; Vouglitois 1983; Wilk et al. 1997; D. Clark, personal communication). Varying numbers have been collected in New Jersey estuaries south of Barnegat Bay; i. e. within Hereford Inlet ( Allen et al. 1978). Although formerly relatively abundant, juvenile scup have not occurred in large numbers in vegetated sites in lower Chesapeake Bay ( Orth and Heck 1980; MAFMC 1996). However, in fall they are still collected in relatively large numbers by the NEFSC trawl surveys at the mouth of the bay ( Figure 12). While juveniles do not occur to any great extent in seaside bays of Maryland and Virginia ( Arve 1960; Schwartz 1961, 1964), Richards and Castagna ( 1970) did find them in their survey of Virginia's seaside bays. The NEFSC groundfish surveys ( 1963 1997) mostly post date the last period of high scup abundance, approximately 1950 1965 ( Northeast Fisheries Science Center 1997). The NEFSC bottom trawl survey results for 1963 1964 ( not shown) indicated that juveniles were widespread and distribution was similar to the present. The only apparent change in this general coastal distribution pattern was in the late 1960s ( during the period of relatively low abundance) when the largest collections of juveniles were clustered off southern New England, Virginia, and North Carolina. This distribution pattern raised the question of whether there were two stocks in the Middle Atlantic Bight ( Hamer 1970).

ADULTS Adults have been reported as far north as the Bay of Fundy, southern Nova Scotia, and Sable Island Bank ( east of Nova Scotia) as summer visitors ( Scott and Scott 1988) and at least as far south as Cape Hatteras. As part of a temperate, migrant guild, scup have even been collected occasionally on the southern Grand Banks ( Brown et al. 1996). Scup occur primarily in the Middle Atlantic Bight. They migrate from offshore winter habitats into coastal waters from Chesapeake Bay to southern New England where they reside from spring to fall ( Bigelow and Schroeder 1953; Richards 1963a; Scott and Scott 1988; Morse 1978; Chang 1990). These migration patterns are reflected in the results of the NEFSC bottom trawl surveys ( Figure 12) and in the Massachusetts inshore survey ( Figure 13). During warm months, larger scup occur in or near the mouths of larger bays, such as Narragansett Bay ( Figures 14, 15) and Long Island Sound ( Figure 16), and along the coast within the 38 m contour ( Morse 1978). Distribution and abundance of adult scup off New England is temperature dependent ( Mayo 1982; Gabriel 1992). Smaller fish are found in more saline (> 15 ppt) shallow bays and parts of estuaries including the Hudson Raritan estuary and Hereford Inlet ( New Jersey) ( Figures 6, 17; Allen et al. 1978; Morse 1978; Werme et al. 1983; Wilk et al. 1997). However, they may not be abundant in all bays; e. g., they have not been reported in Barnegat Bay ( Marcellus 1972; Vouglitois 1983; Tatham et al. 1984), Maryland bays ( MAFMC 1996), or in New York Harbor ( Stoecker et al. 1992; Will and Houston 1992). Adult scup usually arrive offshore in December and winter in deeper water from Nantucket Shoals to Cape Hatteras to depths of about 240 m ( Figures 5 and 12; Pearson 1932; Neville and Talbot 1964; Morse 1978). Scup density and distribution during the winter are related to the location of the 7oC bottom isotherm, their lower preferred limit ( Neville and Talbot 1964). Nesbit and Neville ( 1935) indicated that this band of warmer, outer continental shelf water is influenced mainly by the Gulf Stream just off the shelf. During warm winters, scup can be found across most of the continental shelf south of New Jersey ( Nesbit and Neville 1935). As coastal waters warm above the 7oC threshold in spring, scup return inshore and to the north.

STATUS OF THE STOCKS Commercial landings of scup in the Middle Atlantic Bight have declined substantially since peak landings in the 1950s and early 1960s; although there was a minor peak in landings in the early 1980s ( Figure 18; Northeast Fisheries Science Center 1997). Recreational landings have also declined ( MAFMC 1996). Groundfish surveys by the NEFSC indicated cycles in abundance of scup of about 3 4 years and an overall decline since the 1950 1960s ( Figure 18; Gabriel 1998). Currently, the stock is composed primarily of fish < 3 years old and the age distribution is truncated ( MAFMC 1996). The abundance of scup eggs off southern New England has been low recently ( Gray 1990; Able and Fahay 1998). According to Jeffries and Terceiro ( 1985), slightly warmer average summer temperatures (+ 1 ° C) in coastal waters off southern New England are related to an Page 8 increase in scup abundance. The Middle Atlantic Bight stock is currently considered overfished because the stock is near record low abundance levels and catches exceed Fmax ( Gabriel 1998; National Marine Fisheries Service 1997; Northeast Fisheries Science Center 1997).

RESEARCH NEEDS · The taxonomic status of scup and " southern porgy" should be resolved. · The degree of mixing between populations in the Middle Atlantic and South Atlantic Bights across Cape Hatteras should be determined. · Better characterization of spawning sites and egg and larval habitats is needed. · Offshore winter habitats in the Middle Atlantic Bight need to be identified and described. · The relative importance of larger estuaries ( e. g., Chesapeake, Delaware, and Raritan Bays, Long Island Sound) compared to smaller estuaries and inshore areas ( e. g., Barnegat Bay, seaside bays from Maryland to Virginia) as primary nurseries should be examined. · Determine whether the patchy, inconsistent occurrence of juveniles results from inadequate monitoring or highly variable recruitment. · The habitat factors that result in the patchy distributions of juvenile and adult scup in space and time need to be identified. · The role of natural and artificial structured habitats in the life history, productivity, and fishery management of scup should be determined. · Research should be conducted on the trophic relationships of scup, including the factors that control the production and distribution of their prey ( Kline 1997). · The effects of altering the population age structure on habitat requirements should be examined. · The effects of the winter trawl fishery in the southern Middle Atlantic Bight on spawning stock, juvenile survival, and habitat should be determined. · Information is needed on the direct and indirect effects of degraded environments on feeding, growth, fecundity, survival, and distribution of scup; indirect effects should include food web alterations. · The long term, synergistic effects of combinations of environmental variables ( e. g., pH and toxins) on survival, reproduction, and genetic changes should be investigated ( Kline 1997).

ACKNOWLEDGMENTS C. Steimle, J. Berrien, and R. Ramsey Cross provided publications, literature searches, and interlibrary loans of material. Assistance was also provided by EFH team members ( S. Griesbach, D. Packer, D. Sheehan, W. Morse, D. McMillan, R. Pikanowski, and J. Vitaliano) and others who supplied data for map development and text. W. Gabriel and F. Almeida led us to unpublished information and commented on earlier drafts of this document.

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MAB = Middle Atlantic Bight, SNE = southern New England, GOM = Gulf of Maine.

Life Stage Time of Year Size and Growth Geographic Location Habitat Substrate Temperature Eggs May Aug, south to north progression 0.8 1.0 mm Coastal Virginia  SNE, southern GOM Water column, < 30 m in depth Buoyant in water column 11 23 ° C; most common 12 14 ° C Larvae May Sept, south to north Hatch at ~ 2.0 mm; stage lasts to ~ 15 30 mm MAB and southern GOM, near shore; mostly SNE Water column, < 20 m until juvenile transition In water column until transition 14 22 ° C; peak densities at 15 20 ° C YOY and older juveniles May Nov, south to north YOY: 15 30 mm to 10 cm by Nov; juveniles: to 16 cm by end of 1+ yr MAB GOM; in estuaries spring to fall Estuarine and coastal; from intertidal to about 38 m Sand, mud, mussel, and eel grass beds Greater than ~ 9 27 ° C; mostly 16 22 ° C Winter juveniles Nov Apr/ May ~ 10 13 cm; growth rate reduced Most move offshore and south of New Jersey to warmer, deeper waters; some overwinter in Long Island Sound Mostly > 38 m depth; mid and outer continental shelf; sometime in deep estuaries Poorly known, found over various sand substrates Greater than ~ 7 ° C Summer adults Apr Dec > 15.5 cm FL Coastal from Delaware to GOM ~ 2 38 m Fine to siltysand mud, mussel beds, rock, artificial reefs, wrecks, and other structures ~ 7 25 ° C; can acclimate to 35.6 ° C Winter adults Jan Mar > 15.5 cm FL Most move offshore and south of New Jersey to warmer, deeper waters. Mostly 38 185 m depths; mid/ outer continental shelf. Poorly known, found over various sands. > 7 ° C Spawning adults May Aug, peak in June > 15.5 cm FL; mature at about age 2 Inshore from Delaware Bay north to SNE; mostly in SNE < 30 m, during inshore migration Weedy to sandy > 9 24 ° C Page 15 Table 1. cont'd.

Life Stage Salinity Prey Predators Notes Eggs > 15 ppt Most planktivores where the eggs are found. Eggs hatch in 70 75 hrs at 18 ° C, and in 40 54 hrs at 21 ° C Larvae > 15 ppt Can use yolk for ~ 3 days; at ~ 2.8 mm feeding on zooplankton must begin Most planktivores where the larvae are found. Benthic settlement and juvenile transition occurs at ~ 15 30 mm FL YOY and older juveniles > 15 ppt Small benthic invertebrates, fish eggs and larvae Bluefish, cod, hake, summer flounder, weakfish, striped bass, and others Diurnal schooling feeders. Most migrate to deeper/ warmer waters to the south in winter Winter juveniles Mostly > 30 ppt, except in estuaries Poorly known; possibly small benthic invertebrates, but feeding may be reduced Cod during SNE migration Migrate offshore as temperatures fall below 8 9 ° C and inshore and north as water warms to > 7 ° C; early arrivals can be affected by late cold spell Summer adults > 15 ppt Benthic and near bottom invertebrates, and small fish Sharks, stingrays, dogfish, bluefish, silver hake, black sea bass, and others Usually found in schools of similarly sized individuals. Possibly tolerant or avoid hypoxic conditions Winter adults > 30 ppt Poorly known, but feeding may be reduced Sharks, stingrays, dogfish, bluefish, silver hake, black sea bass, and others 7 ° C isotherm greatly influences distribution Spawning adults > 15 ppt Poorly known, but feeding may be reduced Sharks, stingrays, dogfish, bluefish, silver hake, black sea bass, and others Spawning is often in AM; fish may avoid hypoxic areas Page 16 Figure 1. The scup, Stenotomus chrysops ( from Goode 1884). Page 17 Figure 2. Abundance of the major items in the diet of juvenile ( 1 10 cm) and adult ( 11 40 cm) scup collected during NEFSC bottom trawl surveys from 1973 1980 and 1981 1990. Abundance in the 1973 1980 samples is defined by mean percent prey weights, and in the 1981 1990 samples as mean percent prey volume. The " Arthropoda" are almost entirely crustacea; see text for discussion of specific taxa involved. 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 54.9%

Unknown Animal Remains 18.3% Annelida 11.1% Echinodermata 5.5% Miscellaneous Materials 5.1% All Other Prey 5.0% Echinodermata 52.4%

Arthropoda 20.2% Mollusca 13.1% Unknown Animal Remains 5.6% Annelida 4.1% All Other Prey 4.6%

Annelida 17.0%

Arthropoda 31.1%

Unknown Animal Remains 51.9% Annelida 61.6%

Arthropoda 17.8% Unknown Animal Remains 10.8% All Other Prey 9.8% 1 10 cm ( n= 239) 11 40 cm ( n= 795) a) 1973 1980 1 10 cm ( n= 50) 11 40 cm ( n= 330) b) 1981 1990 Page 18 Figure 3. Abundance of scup eggs relative to water column temperature ( to a maximum of 200 m) and bottom depth from NEFSC MARMAP ichthyoplankton surveys ( May to August 1978 1987, all years combined). Open bars represent the proportion of all stations surveyed, while solid bars represent the proportion of the sum of all standardized catches ( number/ 10 m2). Eggs vs. Temperature Mean Water Column Temperature ( 0 200m, C) 4 6 8 10 12 14 16 18 20 22 24 26 28 0 10 20 50 60 Stations Catch Eggs vs. Depth Depth Interval ( m), Midpoint 10 30 50 70 90110130150 170190 210230250 270290 325375450 750 12501750> 2000 Percent 0 10 20 90 100 Scup Eggs, May to August Page 19 Figure 4. Abundance of scup larvae relative to water column temperature ( to a maximum of 200 m) and bottom depth from NEFSC MARMAP ichthyoplankton surveys ( July and August 1977 1987, all years combined). Open bars represent the proportion of all stations surveyed, while solid bars represent the proportion of the sum of all standardized catches ( number/ 10 m2). Larvae vs. Temperature Mean Water Column Temperature ( 0 200m, C) 4 6 8 10 12 14 16 18 20 22 24 26 28 0 10 20 90 100 Stations Catch Larvae vs. Depth Depth Interval ( m), Midpoint 10 30 50 70 90110130150170190210230250 27029032537545075012501750> 2000 Percent 0 10 20 30 90 100 Scup Larvae, July & August Page 20 Figure 5. Seasonal abundance of juvenile and adult scup relative to bottom water temperature and depth based on NEFSC bottom trawl surveys ( 1963 1997, all years combined). Open bars represent the proportion of all stations surveyed, while solid bars represent the proportion of the sum of all standardized catches ( number/ 10 m2). SPRING 0 5 10 15 20 25 30 35 4 6 8 10 12 14 16 18 20 BOTTOM TEMPERATURE ( C) PERCENT STATIONS N= 643 CATCHES N= 106979 SUMMER 0 5 10 15 20 25 30 35 7 9 11 12 13 14 15 16 17 18 19 20 21 22 25 BOTTOM TEMPERATURE ( C) PERCENT STATIONS N= 109 CATCHES N= 14653 FALL 0 2 4 6 8 10 12 14 16 7 9 11 13 15 17 19 21 23 25 27 BOTTOM TEMPERATURE ( C) PERCENT STATIONS N= 1879 CATCHES N= 384403 WINTER 0 5 10 15 20 25 30 35 40 45 4 5 6 7 8 9 10 11 12 13 BOTTOM TERATURE ( C) PERCENT STATIONS N= 133 CATCHES N= 31625 SPRING 0 5 10 15 20 25 30 35 0 20 40 60 80 100 120 140 160 180 200 220 270 DEPTH ( m) PERCENT STATIONS N= 643 CATCHES N= 106979 SUMMER 0 10 20 30 40 50 60 70 80 0 10 20 30 40 50 DEPTH ( m) PERCENT STATIONS N= 109 CATCHES N= 18508 FALL 0 5 10 15 20 25 30 35 40 45 50 0 20 40 60 80 100 160 210 DEPTH ( m) PERCENT STATIONS N= 1879 CATCHES N= 384404 WINTER 0 5 10 15 20 25 30 35 20 40 60 80 100 120 140 310 DEPTH ( m) PERCENT STATIONS N= 133 CATCHES N= 31625 Juveniles: £ 15 cm TL BOTTOM TEMPERATURE ( C) Page 21 Figure 5. cont'd. SPRING 0 5 10 15 20 25 30 35 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 BOTTOM TEMPERATURE ( C) PERCENT STATIONS N= 643 CATCHES N= 42311 SUMMER 0 5 10 15 20 25 30 35 40 7 9 11 12 13 14 15 16 17 18 19 20 21 22 25 BOTTOM TEMPERATURE ( C) PERCENT STATIONS N= 109 CATCHES N= 3855 FALL 0 2 4 6 8 10 12 14 16 18 7 9 11 13 15 17 19 21 23 25 27 BOTTOM TEMPERATURE ( C) PERCENT STATIONS N= 1879 CATCHES N= 49202 WINTER 0 10 20 30 40 50 60 70 80 4 5 6 7 8 9 10 11 12 13 BOTTOM TEMPERATURE ( C) PERCENT STATIONS N= 133 CATCHES N= 7063 SPRING 0 2 4 6 8 10 12 14 16 18 20 0 20 40 60 80 100 120 140 160 180 200 220 270 DEPTH ( m) PERCENT STATIONS N= 643 CATCHES N= 42311 SUMMER 0 10 20 30 40 50 60 70 0 10 20 30 40 50 70 DEPTH ( m) PERCENT STATIONS N= 109 CATCHES N= 3855 FALL 0 5 10 15 20 25 30 35 40 45 50 0 20 40 60 80 100 160 210 DEPTH ( m) PERCENT STATIONS N= 1879 CATCHES N= 49202 WINTER 0 10 20 30 40 50 60 20 40 60 80 100 120 140 310 DEPTH ( m) PERCENT STATIONS N= 133 CATCHES N= 7063 Adults: > 15 cm TL Page 22 Figure 6. Abundance of juvenile and adult scup relative to bottom water temperature, depth, dissolved oxygen, and salinity based on Hudson Raritan estuary trawl surveys ( 1992   1997, all years combined). Temperature ( C)

Depth ( ft) 0 2 4 6 8 10 12 14 16 18 20 22 24 26 0 5 10 15 20 25 Stations Catches 10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85 0 5 10 15 20 25 30 35 Dissolved Oxygen ( mg/ l)

Salinity ( ppt) 0 1 2 3 4 5 6 7 8 9 10 11 12 13 0 5 10 15 20 25 30 35 40 45 15 17 19 21 23 25 27 29 31 33 35 0 2 4 6 8 10 12 14 16 18 Adults (> 15 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 18 20 Stations Catches 10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85 0 5 10 15 20 25 30 35 Dissolved Oxygen ( mg/ l)

Salinity ( ppt) 0 1 2 3 4 5 6 7 8 9 10 11 12 13 0 5 10 15 20 25 30 35 15 17 19 21 23 25 27 29 31 33 35 0 2 4 6 8 10 12 14 16 Juveniles ( # 15 cm) Page 23 Figure 7. Abundance of juvenile and adult scup relative to bottom water temperature and depth based on Massachusetts inshore bottom trawl surveys ( spring and autumn 1978 1996, all years combined). Open bars represent the proportion of all stations surveyed, while solid bars represent the proportion of the sum of all standardized catches ( number/ 10 m2). 1 3 5 7 9 11 13 15 17 19 21 23 0 10 20 30 40 50 1 3 5 7 9 11 13 15 17 19 21 23 0 5 10 15 20 25 0 10 20 30 40 50 0 10 20 30 40 0 10 20 30 40 0 10 20 30 40 1 3 5 7 9 11 13 15 17 19 21 23 0 4 8 12 16 20 1 3 5 7 9 11 13 15 17 19 21 23 0 10 20 30 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 Scup Page 24 Figure 8. Seasonal abundance of juvenile and adult scup relative to mean bottom water temperature and bottom depth from Rhode Island Narragansett Bay trawl surveys, 1990 1996. Open bars represent the proportion of all stations surveyed, while solid bars represent the proportion of the sum of all catches.  1 1 3 5 7 9 11 13 15 17 19 21 23 25 27 0 10 20 30  1 1 3 5 7 9 11 13 15 17 19 21 23 25 27 0 10 20 30 40  1 1 3 5 7 9 11 13 15 17 19 21 23 25 27 0 10 20 30 40  1 1 3 5 7 9 11 13 15 17 19 21 23 25 27 0 20 40 60 80 Bottom Temperature ( C) Winter Spring Summer Autumn Scup Juveniles (< 16 cm)

Stations Catches 10 20 30 40 50 60 70 80 90 100 110 120 0 20 40 60 80 10 20 30 40 50 60 70 80 90 100 110 120 0 10 20 30 40 50 10 20 30 40 50 60 70 80 90 100 110 120 0 20 40 60 10 20 30 40 50 60 70 80 90 100 110 120 0 10 20 30 Bottom Depth ( ft) Winter Spring Summer Autumn Scup Juveniles (< 16 cm)

Stations Catches Page 25 Figure 8. cont'd.  1 1 3 5 7 9 11 13 15 17 19 21 23 25 27 0 10 20 30 40 50  1 1 3 5 7 9 11 13 15 17 19 21 23 25 27 0 10 20 30  1 1 3 5 7 9 11 13 15 17 19 21 23 25 27 0 10 20 30 40  1 1 3 5 7 9 11 13 15 17 19 21 23 25 27 0 4 8 12 16 20 Bottom Temperature ( C) Winter Spring Summer Autumn Scup Adults ( $ 16 cm) Stations Catches 10 20 30 40 50 60 70 80 90 100 110 120 0 10 20 30 10 20 30 40 50 60 70 80 90 100 110 120 0 10 20 30 40 50 10 20 30 40 50 60 70 80 90 100 110 120 0 20 40 60 10 20 30 40 50 60 70 80 90 100 110 120 0 10 20 30 40 50 Bottom Depth ( ft) Scup Adults ( $ 16 cm) Stations Catches Winter Summer Autumn Spring Page 26 Figure 9. The distribution of scup from Newfoundland to Cape Hatteras. Data are from the U. S. NOAA/ Canada DFO East Coast of North America Strategic Assessment Project ( http//: www orca. nos. noaa. gov/ projects/ ecnasap/ ecnasap_ table1. html). Page 27 Figure 10. Distribution and abundance of scup eggs collected during NEFSC MARMAP ichthyoplankton surveys, 1978 1987 [ see Reid et al. ( 1999) for details]. The upper left figure is a summary of all months and years; the remaining figures are by individual month ( May, June, July and August) for all years combined. Scup ( Stenotomus chrysops)

Eggs MARMAP Ichthyoplankton Surveys 61 cm Bongo Net; 0.505 mm mesh 1978 to 1987 ( May, Jun, Jul, Aug)

Number of Tows = 3438; with eggs = 12 76 75 74 73 72 71 70 69 68 67 66 65 35 36 37 38 39 40 41 42 43 44 45 Eggs / 10m2 1 to < 10 10 to < 100 100 to 149 76 75 74 73 72 71 70 69 68 67 66 65 35 36 37 38 39 40 41 42 43 44 45 Scup ( Stenotomus chrysops)

Eggs MARMAP Ichthyoplankton Surveys 61 cm Bongo Net; 0.505 mm mesh May, 1978 to 1987 Number of Tows = 1085; with eggs = 1 Monthly Mean Density = 0.003 Eggs/ 10m2 Eggs / 10m2 None 1 to 3 76 75 74 73 72 71 70 69 68 67 66 65 35 36 37 38 39 40 41 42 43 44 45 Scup ( Stenotomus chrysops)

Eggs MARMAP Ichthyoplankton Surveys 61 cm Bongo Net; 0.505 mm mesh June, 1978 to 1987 Number of Tows = 709; with eggs = 3 Monthly Mean Density = 0.240 Eggs/ 10m2 Eggs / 10m2 None 1 to < 10 10 to < 100 100 to 149 76 75 74 73 72 71 70 69 68 67 66 65 35 36 37 38 39 40 41 42 43 44 45 Scup ( Stenotomus chrysops)

Eggs MARMAP Ichthyoplankton Surveys 61 cm Bongo Net; 0.505 mm mesh July, 1978 to 1987 Number of Tows = 781; with eggs = 7 Monthly Mean Density = 0.161 Eggs/ 10m2 Eggs / 10m2 None 1 to < 10 10 to 59 Page 28 Figure 10. cont'd. 76 75 74 73 72 71 70 69 68 67 66 65 35 36 37 38 39 40 41 42 43 44 45 Scup ( Stenotomus chrysops)

Eggs MARMAP Ichthyoplankton Surveys 61 cm Bongo Net; 0.505 mm mesh August, 1978 to 1987 Number of Tows = 863; with eggs = 1 Monthly Mean Density = 0.008 Eggs/ 10m2 Eggs / 10m2 None 1 to 7 Page 29 Figure 11. Distribution and abundance of scup larvae collected during NEFSC MARMAP ichthyoplankton surveys, 1977 1987 [ see Reid et al. ( 1999) for details]. The upper left figure is a summary of all months and years; the remaining figures are by individual month ( July and August) for all years combined. 76 75 74 73 72 71 70 69 68 67 66 65 35 36 37 38 39 40 41 42 43 44 45 Scup ( Stenotomus chrysops)

Larvae MARMAP Ichthyoplankton Surveys 61 cm Bongo Net; 0.505 mm mesh 1977 to 1987 ( Jul & Aug)

Number of Tows = 2086; with larvae = 2 Larvae / 10m2 1 to < 10 10 to 98 76 75 74 73 72 71 70 69 68 67 66 65 35 36 37 38 39 40 41 42 43 44 45 Scup ( Stenotomus chrysops)

Larvae MARMAP Ichthyoplankton Surveys 61 cm Bongo Net; 0.505 mm mesh July, 1977 to 1987 Number of Tows = 938; with larvae = 1 Monthly Mean Density = 0.104 Larvae / 10m2 Larvae / 10m2 None 1 to < 10 10 to 98 76 75 74 73 72 71 70 69 68 67 66 65 35 36 37 38 39 40 41 42 43 44 45 Scup ( Stenotomus chrysops)

Larvae MARMAP Ichthyoplankton Surveys 61 cm Bongo Net; 0.505 mm mesh August, 1977 to 1987 Number of Tows = 1148; with larvae = 1 Monthly Mean Density = 0.003 Larvae / 10m2 Larvae / 10m2 None 1 to 4 Page 30 Figure 12. Distribution and abundance of juvenile and adult scup collected during NEFSC bottom trawl surveys ( 1963 1997, all years combined). 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 31 Figure 12. cont'd. Page 32 Figure 13. Distribution and abundance of juvenile and adult scup in Massachusetts coastal waters collected during spring and autumn Massachusetts inshore bottom trawl surveys, 1978 1996 [ see Reid et al. ( 1999) for details]. Scup Mass. Inshore Trawl Survey Spring 1978  1996 Juveniles (< 16cm)

Number/ Tow 1 to 10 10 to 50 50 to 250 250 to 1000 1000 to 5233 Scup Mass. Inshore Trawl Survey Autumn 1978  1996 Juveniles (< 16cm)

Number/ Tow 1 to 50 50 to 100 100 to 500 500 to 1000 1000 to 5233 Scup Mass. Inshore Trawl Survey Spring 1978  1996 Adults (>= 16cm)

Number/ Tow 1 to 25 25 to 100 100 to 500 500 to 1000 1000 to 6872 Scup Mass. Inshore Trawl Survey Autumn 1978  1996 Adults (>= 16cm)

Number/ Tow 1 to 10 10 to 50 50 to 100 100 to 500 500 to 1817 Page 33 Figure 14. Seasonal distribution and abundance of juvenile and adult scup collected in Narragansett Bay during 1990 1996 Rhode Island bottom trawl surveys. The numbers shown at each station are the average catch per tow rounded to one decimal place [ see Reid et al. ( 1999) for details]. 0.0 0.0 0.0 0.0 0.1 0.0 0.0 0.0 0.0 0.0 0.1 0.0 0.0 7.4 2.8 0.0 0.1 2.7 0.1 0.0 0.2 6.0 9.1 16.5 0.3 5.8 9.8 37.3 9.1 3.2 99.1 65.5 105.8 17.1 537.2 172.7 16.8 112.5 18.8 318.6 56.9 37.2 25.2 39.5 171.8 66.8 113.2 210.7 443.2 124.9 182.9 18.4 Spring Summer Autumn Winter Juveniles ( < 16 cm) Page 34 Figure 14. cont'd. 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 3.0 2.1 0.0 0.0 5.0 0.0 0.3 0.2 23.9 49.8 39.8 0.2 33.4 0.3 7.1 0.0 0.0 13.5 0.0 5.3 1.6 66.6 26.8 2.9 2.3 1.8 7.3 40.8 0.6 1.3 37.1 2.9 0.9 1.1 16.6 4.2 4.6 4.2 1.8 Spring Summer Autumn Winter Adults (>= 16 cm) Page 35 Figure 15. Size frequency distribution of scup collected in Narragansett Bay during 1990 1996 Rhode Island bottom trawl surveys. 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 0 1 2 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 0 200 400 600 800 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 0 1000 2000 3000 4000 5000 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 0 2000 4000 6000 8000 10000 Total Length ( cm) Winter Spring Summer Autumn Page 36 Figure 16. Distribution, abundance, and size frequency of scup in Long Island Sound in spring and autumn, from the Connecticut bottom trawl surveys, 1992 1997 [ see Reid et al. ( 1999) for details]. Number/ Tow 1 to < 50 50 to < 100 100 to < 500 500 to < 1000 1000 to < 2048 Number/ Tow 1 to < 200 200 to < 500 500 to < 1000 1000 to < 2000 2000 to < 2428 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 Scup Scup Page 37 Figure 17. Seasonal distribution and abundance of juvenile and adult scup in the Hudson Raritan estuary collected during Hudson Raritan estuary trawl surveys, 1992   1997 [ see Reid et al. ( 1999) for details]. Staten Island NEW YORK NEW JERSEY No/ Tow 1  9 10  24 25  49 50  199 200  685 Hudson Raritan Estuary Fall 1992  1996 Juveniles (< 16 cm) Scup Staten Island NEW YORK NEW JERSEY No/ Tow 1  9 10  24 25  49 50  199 200  685 No Catches Hudson Raritan Estuary Winter 1992  1997 Juveniles (< 16 cm) Scup Staten Island NEW YORK NEW JERSEY No/ Tow 1  9 10  24 25  49 50  199 200  685 Hudson Raritan Estuary Juveniles (< 16 cm) Scup Spring 1992  1997 Staten Island NEW YORK NEW JERSEY No/ Tow 1  9 10  24 25  49 50  199 200  685 Scup Hudson Raritan Estuary Summer 1992  1996 Juveniles (< 16 cm) Page 38 Figure 17. cont'd. Staten Island NEW YORK NEW JERSEY No/ Tow 1  9 10  24 25  49 50  199 200  685 Hudson Raritan Estuary Fall 1992  1996 Adults (> 15 cm) Scup Staten Island NEW YORK NEW JERSEY No/ Tow 1  9 10  24 25  49 50  199 200  685 Scup Hudson Raritan Estuary Summer 1992  1996 Adults (> 15 cm) Staten Island NEW YORK NEW JERSEY No/ Tow 1  9 10  24 25  49 50  199 200  685 Scup Hudson Raritan Estuary Winter 1992  1997 Adults (> 15 cm)

No Catches Staten Island NEW YORK NEW JERSEY No/ Tow 1  9 10  24 25  49 50  199 200  685 Scup Hudson Raritan Estuary Spring 1992  1997 Adults (> 15 cm) Page 39 Figure 18. Commercial landings ( metric tons, mt) and NEFSC bottom trawl survey indices ( stratified mean catch per tow, kg) for scup in southern New England and the Middle Atlantic Bight. Southern New England and Middle Atlantic Year 1965 1970 1975 1980 1985 1990 1995 2000 Landings ( mt x 1000)

0 2 4 6 8 10 Stratified mean catch/ tow ( kg)

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