[From the U.S. Government Printing Office, www.gpo.gov]





                                                                                                   Science for Solutions



                                                                                                                                                                                                      of"Alf OF
                            NOAA COASTAL OCEAN PROGRAM                                                                                                                                            J,
                            Decision Analysis Series No. 1
                                                                                                                                                                                                        Q@


                            SYNTHESIS OF SUMMER FLOUNDER
                                                                    HABITAT PARAMETERS


                                                                                                     Kenneth W. Able
                                                                                                      Susan C. Kaiser


                                                                                                                May 1994













                                                                                                    11%            SPRING SPIMAIVIER
                                                                                      .........                                 0.


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                                                                                                                                       A





                                                                                   U.S. DEPARTMENT OF COMMERCE
            QL637                                                 National Oceanic and Atmospheric Administration
             .P5A25                                                                                   coastal Ocean Office
            1994










                                  The Decision Analysis Series has been
                                  established by NOAA's Coastal Ocean
                                  Program (COP) to present documents for
                                  coastal resource decision makers which
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               Cover illustration: Life History of Summer Flounder
               (larger view explained in detail on inside back cover)









                                           Science for Solutions


                                                                                         
             NOAA COASTAL OCEAN PROGRAM
             Decision Analysis Series No. 1

                                                                                         


                SYNTHESIS OF SUMMER FLOUNDER
                              HABITAT PARAMETERS





                                            Kenneth W. Able
                                            Susan C. Kaiser



                                          Marine Field Station
                                           Rutgers University
                             Institute of Marine and Coastal Sciences
                                       800 Great Bay Boulevard
                                              Tuckerton, NJ


                                                May 1994




                                    U.S. DEPARTMENT OF COMMERCE
                                           Ronald H. Brown, Secretary
                              National Oceanic and Atmospheric Administration
                                         D. James Baker, Under Secretary
                                            Coastal Ocean Office
                                             Donald Scavia, Director

          
          
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                                                 NOAA/CCEH
                                              1990 HOBSON AVE.
                                              CHAS SC 29408-2623






















                   This Publication should be cited as:

                   Able, Kenneth W and Susan Q. Kaiser.            1994. Synthesis of Summer Flounder Habitat Parameters. NOAA
                   Coastal:Ocean Program Decision Analysis Series No.. 1. NOAA Coastal Ocean Office, Silver Springi MD.
                   68 pp. + biblio. + 3 apps.











                                      This publication does,not constitute an endorsement of any commercial product,or
                                      intend to be an opinion beyond scientifi.c or other results obtained by the National
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                Note to Readers


             -,The NOAA Coastal Ocean Program (COP) provides a focal point through which the agency,
                together with other organizations with responsibilities for the coastal environment and its
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                A specific objective of COP is to provide the highest quality scientific information to coastal
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                series through the use of a panel of multidisciplinary technical experts; 2) to solicit proposals to
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                Seven topics and associate'd principal investigators were selected in the initial round. This
                volume, Synthesis of Summer Flounder Habitat Parameters by Kenneth W. Able and Susan C.
                Kaiser of Rutgers University,,is the first document.in this Decision. Analysis Series to be
                published. Other volumes will be published over the next two years on the following topics:
                seagrass restoration technology, salt marsh restoration, coastal watershed restoration, restoring
                streams and anadromous fish habitat affected by logging, eutrophication and phytoplankton
                blooms, and management of cumulative coastal environmental impacts.

                As with all of its products, COP is very interested in ascertaining the utility of the Decision
                Analysis Series particularly in regard to its application to the management decision process.
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                cover. My Internet address is [email protected].




                                                         /Donald Scavia
                                                            Director
                                                            NOAA Coastal Ocean Program










            Table of Contents


            LIST OF FIGURES AND TABLES                                                  v

            ACKNOWLEDGMENTS                                                            ix


            EXECUTIVE SUMMARY                                                          A


            CHAPTER LINTRODUCTION
                   Background and Overview
                   Scope                                                                2

            CHAPTER 2: DISTRIBUTION AND LIFE HISTORY PATTERNS                           3
                   Geographical Distribution                                            3
                   General Life History                                                 3
                   Stock Identification                                                17


            CHAPTER 3: EGGS                                                            19
                   Introduction                                                        19
                   Spatial and Temporal Distribution                                   19
                   Temperature                                                         19

            CHAPTER 4: OFFSHORE LARVAE                                                 21
                   Introduction                                                        21
                   Spatial and Temporal Distribution                                   21
                   Temperature                                                         19
                   .Substrate                                                          22

            CHAPTER 5: ESTUARINE LARVAE                                                23
                   Introduction                                                        23
                   Spatial and Temporal Distribution                                   23
                   Temperature and Salinity                                            26
                   Substrate and Behavior                                              26








            iv
                                                                        Table of Contents



            CHAPTER 6: ESTUARINE JUVENILES                                           29
                  Introduction                                                       29
                  Spatial and Temporal Distribution..                                29
                  Temperature, Salinity and Dissolved Oxygen                         40
                  Substrate                                                          45
                  Behavior                                                           46


            CHAPTER 7: OFFSHORE JUVENILES                                            47
                  Introduction                                                       47
                  Spatial and Temporal Distribution                                  47
                  Temperature                                                        51

            CHAPTER 8: ESTUARINE AND. OFFSHORE ADULTS                                53
                  Introduction                                                       53
                  Spatial and Temporal Distribution                                  53
                  Temperature, Salinity and Dissolved Oxygen                         58
                  Substrate                                                          59


            CHAPTER-9: IMPLICATIONS FOR RESOURCE MANAGEMENT                          61

            GLOSSARY                                                                 63


            APPENDICES

                  Appendix A.
                  Comprehensive. Summer, Flounder Bibliography                        A-i
                  Author Index                                                        A-51
                  Subject Index                                                       A-59

                  Appendix B.
                  List of Experts                                                     B-1

                  Appendix C.
                  User Groups                                                         C-1









            List of FigureS-and Tables

                                                Figures

            2.1   East coas.t of North America location maps showing summer flounder range.
            2.2   Cumulative egg distribution and abundance over northern and mid-Atlantic
                  continental shelf.

            2.3   Egg abundance'by Month and r6gio n'6ver the'' north and mid-Atlantic
                  continental shelf.

            2.4   Monthly larval distribution and abundance'over, the -north and- mid-Atlantic
                  continental shelf.

            2.5   Larval abundance by month and region over ther north and mid-Atlantic
                  continental shelf.'

            2.6   Transformation stages of summer flounder.

            2.7   Locations of summer flounder studies: (A) Great Bay-Little -Egg Harbor, New
                  Jersey; (6) Charleston Harbor, South Carolina; (C) Long Island Sound, New
                  York/Connecticut; and (D) Pamlico-Albemarle sounds, North Carolina..

            2.8  -Cumulative length- and stage:-frequency distributions for transformi ng -larvae in
                  two marsh creeks in Great Bay-Little Egg Harbor.

            2.9   Monthly length-frequency distributions for juveniles in Great Bay-Little Egg
                  Harbor.


            2.10  Seasonal distribution on the north and mid-Atlantic continental shelf.

            2.11  Synopsis of seasonal movements from tagging studies.

            5.1   Seasonal variation in abundance of transforming larvae in Little Sheepshead
                  Creek, Great Bay-Little Egg Harbor..








             Vi
                                                                               Ficiures and Tables


             5.2    Seasonal variation in abundance of -transforming larvae in Schooner Creek,
                    Great Bay-Little Egg Harbor.

             6.1    Monthly length-frequency'distri'b'utions for juveniles in four marsh creeks in
                    Charleston Harbor.

             6.2    Monthly  length-frequency distributions of juveniles and adults in Long Island
                    Sound.

             6.3    Monthly distribution and abundance of juveniles and adults in Long Island
                    Sound.


             6.4    Seasonal length-frequency distributions for juveniles and adults in Pamlico-
                    Albemarle sounds.

             6.5    Monthly length-frequency distributions for juveniles in shallow marsh habitats in
                    Pamlico-Albemarle sounds.

             6.6    Seasonal distribution and abundance of juveniles and adults in Pamlico-
                    Albemarle sounds.


             6.7    Bimonthly length-frequency distributions for juveniles in Charleston Harbor.

             6.8    Juvenile abundance relative to salinity in four marsh creeks in Charleston
                    Harbor.


             6.9    Habitat-specific growth of caged juveniles in Great Bay-Little Egg Harbor.

             7.1    Seasonal distribution of juveniles on the north and mid-Atlantic continental
                    shelf.


             7.2    Sampling locations for two inner continental shelf trawl surveys in the South
                    Atlantic Bight.

             7.3    Monthly length-frequency distributions for juveniles and adults on the South
                    Atlantic Bight inner continental shelf.

             8.1    Monthly distribution of adults on the north and mid-Atlantic continental shelf
                    from the commercial fishery.

             8.2    July length-frequency distribution s of adults in Great Bay from the recreational
                    fishery over a 10-year period.






              Figures and Tables
                                                                                                     Vii



                                                       Table-s,


              6.1    Characteristics of marsh, creek habitat. for juveniles in Charleston Harbor.

              6.2    Characteristics of marsh creek habitat for juveniles in Chesapeake Bay.































Ad









              Acknowledgments

              Numerous individuals and agencies contributed their time and/or resources toward the
              completion of this project. Special thanks go to the following individuals who provided
              unpublished data sets and base maps used extensively in this synthesis: M. Fahay
              (National Marine Fisheries Service, Sandy Hook Laboratory) for offshore egg and
              larval distribution data; T. Azarovitz, G. Shepherd, M. Terceiro and B. O'Gorman
              (National Marine Fisheries Service, Woods Hole Laboratory) for offshore adult
              distribution data; P. Howell and M. Johnson (Connecticut Bureau of Natural Resources
              - Marine Fisheries) for juvenile and adult distribution and length data; R. Jesien and C.
              Hocutt (University of Maryland, Horn Point Environmental Laboratory) for tagging data;
              L. Daniel III (Virginia Institute of Marine Sciences) for habitat data on juveniles in
              marsh creeks; J. Musick and J. Desfosse (Virginia Institute of Marine Sciences) for
              tagging data; L. Mercer and J.P. Monaghan Jr. (North Carolina Division of Marine
              Fisheries) for juvenile and adult distribution, length and tagging data; C. Wenner
              (South Carolina Marine Resources Center) for offshore South Atlantic@ Bight adult
              distribution and length data and habitat data on juveniles in marsh creeks.

              The following individuals provided unpublished data that supported our work: A.
              Burnett and D. Byrne (New Jersey Div   ision of Fish, Game and Wildlife); P-Carlsen
              (American Littoral Society); J. Cagey (Maryland Department of Natural Resources); M.
              Fonseca (National Marine Fisheries Service, Beaufort Laboratory); T. Lynch (Rhode
              Island Division of Fish and Wildlife); K. McKown (New York Department of
              Environmental Conservation); S. Michels (Delaware Division of Fish and Wildlife); C.
              Powell (University of Rhode Island); C. Epifanio, K. Malloy and T. Targett (University of
              Delaware).

              Our literature search was facilitated by unpublished bibliographies contributed by: T.
              Azarovitz and R. Rountree (National Marine Fisheries Service, Woods Hole
              Laboratory); D. Packer and A. Studholme (National Marine Fisheries Service, Sandy
              Hook Laboratory); R. Matheson (Florida Division of Natural Resources); J. Monaghan
              Jr. (North Carolina Division of Marine Fisheries).

              The synthesis benefitted from discussions with and literature received from many
              individuals, including: J. Burke and D. Peters (National Marine Fisheries Service,
              Beaufort Laboratory); W. Castonguay (Massachusetts Division of Marine Fisheries); G.
              Gilm'ore (Harbor Branch Oceanographic Institute); T. Hoff (Mid-Atlantic Marine
              Fisheries Council); D. Hoss (National Marine Fisheries Service, Beaufort Laboratory);











             x
                                                                             Acknowledgements


             J. Pafford and S. Shipman (Georgia Department of Natural Resources); A. Powell and
             F. Schwartz (University of North Carolina)..

             L. Markoff and T. Turrin, Rutgers University, Nelson. Biological Laboratories Computer
             Lab, provided graphics support.

             The following Rutgers University Marine Field Station personnel helped prepare the
             synthesis: E. Bender and E. Duval assisted with data entry and manipulation; A.
             Tonkin entered bibliographic data; B.r Zlotnik provided,word processing and editorial
             assistance. We sincerely thank all of the above and apologize to anyone inadvertently
             omitted. This work was supported by the National Oceanic and Atmospheric
             Administration Coastal Ocean@Peogram, and Rutgers University, Institute of Marine and
             Coastal Sciences' (IMCS).  This i's IMCS Contribution Number 94-14.









              Executive SumMary


              The summer flounder, Paralichthys dentatus is overexploited, and. is currently at very
              low levels of abundance. This is reflectedIin the compressed 'age structure of the.
              population and the low catches in both commercial, and recreational fisheries.
              Declining habitat quantity and quality,pay,be.-contrO, uting.tothes.e declines,,,h.owe.ver
              we lack a thorougo understandi,ng of the role of.habitats in the. population dynamics of
              this species. Stock structure is unresolved and current interpretations, depending on
              the technique and study area, suggest that there may be two or three spawning
              populations. If so, these stocks may have differing habitat requirements. In response
              to this lack of knowledge, this document summarizes and synthesizes the available
              information on summer flounder habitat in all life history stages (eggs, larvae, juveniles
              and adults) and identifies areas where further research is needed.

              Several levels of investigation were conducted in order to produce this document.
              First, an extensive search for summer flounder habitat information was made, which
              included both the primary and gray literature as well as unanalyzed data. Second,
              state and federal fisheries biologists and resource managers in all states within the
              primary range of summer flounder (Massachusetts to Florida) were interviewed along
              with a number of fish ecologists and summer flounder experts from the academic and
              private sectors. Finally, information from all sources was analyzed and synthesized to
              form a coherent overview.


              This document first presents an overview of the economic importance and current
              status of summer flounder (Chapter 1). It then summarizes our present state of
              knowledge of summer flounder distribution, life history patterns and stock identification
              (Chapter 2). This is followed by a synopsis of habitat requirements during each life
              history stage. For convenience, this is presented by general habitat as offshore eggs
              (Chapter 3), offshore larvae (Chapter 4), estuarine larvae (Chapter 5), estuarine
              juveniles (Chapter 6), offshore juveniles (Chapter 7) and estuarine and offshore adults
              (Chapter 8). In several instances, previously undigested data sets are analyzed to
              provide more detailed information, especially for estuarine juveniles. The information
              is then discussed in terms of its relevance to resource managers (Chapter 9).

              A comprehensive bibliography on all aspects of the distribution, biology, and ecology
              of summer flounder (Appendix A) is provided with both an author index and a subject
              index for easy reference. This bibliography also serves as the primary reference for











             X11
                                                                                Executive Summa!y


             literature cited in the text. Finally, a list of researchers who are considered experts on
             summer flounder (Appendix B) and a list of potential user groups of this document
             (Appendix C) are included.

             Summer flounder occur in    continental shelf and estuarine waters of the western    north
             Atlantic from Nova Scotia (Canada) to Florida (United States). The center of
             abundance lies in the Middle Atlantic Bight, and due to the extensive fishery and
             research emphasis, our understanding of'life history patterns is most complete for this
             portion of the range. In the Middle Atlantic Bight,-temporal and spatial distribution in
             estuarine and offshore habitats is determined largely by temperature and salinity.
             Generally, adult summer flounder and older juveniles migrate seasonally in response
             to temperature changes, spending winters -on the middle and outer continental shelf
             and summers on the inner continental shelf and in estuaries..' Adults spawn while
             moving offshore in autumn and early winter. Eggs rise to near surface waters and the
             newly hatched larvae are planktonic and symmetrical in shape. While over the
             continental shelf, larvae begin transformation, the process. by which the right eye
             migrates to the left side of the head. This is accompanied by other morphological and
             physiological changes as larvae prepare -for settlement. During winter to early spring,
             transforming larvae move into estuaries where eye migration is com     Ipleted and
             settlement to the bottom marks the beginning    .of the juvenile stage. In spring, many
             fishes that have overwintered on the continental shelf join these juveniles in estuaries.
             During their summer residencies, juveniles and adults are most abundant in higher -
             salinity waters of estuaries. As winter approaches, most juveniles move offshore with'
             adults, however some may overwinter in the deep waters of larger estuarine systems.
             In the South Atlantic Bight, summer flounder life history patterns have not been as
             thoroughly studied. In general, adults spawn on the continental shelf in autumn and
             winter, and transforming larvae enter estuarine nursery habitats in the spring where
             they settle. However, adults may predominantly use inner continental shelf waters as
             summer forage grounds rather than estuaries.

             This synt ,hesis indicates that temperature and dissolved oxygen are the habitat
             parameters of primary importance to summer flounder. Low winter temperatures are
             likely a source of natural mortality to transforming larvae as they enter estuaries,
             especially in the northern portion of their range. Low spring temperatures decrease
             growth rates for transforming larvae and early juveniles, and by delaying or slowing
             growth, may make these individuals more vulnerable to predation for longer periods of
             time, thus reducing survival. The presumed preference for higher salinity explains the
             greater abundance in the lower portions of estuaries. Low dissolved oxygen on the
             continental shelf and in estuaries can affect distribution and survivorship. Episodes of
             hypoxia or anoxia can cause habitat use patterns to change as individuals attempt to
             migrate away from feeding areas. These movements may concentrate individuals in a
             small area, making them more susceptible to fishing mortality. If migration is not






               Executive Summa[y
                                                                                                       Xiii



               possible, natural mortality can result, which together with fishing losses can have a
               significant- impact on local populations.

               In summary, despite the economic importance of summer flounderin fisheries along.,
               the east coast of the U.S., we still lack a clear understanding.of habitat requirements
               and, this, is especially true for the.eggs and larvae. We know somewhat more for the.
               juve niles and adults, although the depth of understanding varies from region. to region.
               Due to the highly migratory naty.reof this species, summer flounder exploit a variety of
               habitats from shallow estuaries to the deep. edge of,the continental shelf., However,
               the habitats that are perhaps the most critical, estuarine nurseries, are the habitats.
               that are most severely impacted by human activity.

               As a result, we recommendthat estuarine'jyveniles should     be th eJocus.of researchers
               and resource managers because: 1) juvenile growthand survival in estuarine nurseries
               may be especially critical to subsequent year-class strength; and 2) estuarine habitats
               are especially vulnerable to alteration and negative impacts that could.influence habitat
               quantity and quality. - Habitat-specific data is generally limited. Of those habitats
               examined, high-salinity subtidal salt Mars h..cree ks and-sha,ilow portions of bays
               appear to be most important as nurseries, especially -for.the early.j'uvenile stages.. We
               therefore suggest-that resource managers    pay particular attention to these habitats
               and their maintenance in order to  improve the status of su mmer flounder populations..










             Chapter I
             Introduction


             BACKGROUND AND OVERVIEW


             The summer flounder, (Paralichthys dentatus , an important species in commercial and
             recreational fisheries on the east coast of the United States, is currently overexploited
             (Mid-Atlantic Fishery Management Council 1990, 1991a,b). Since implementation of
             the summer flounder management plan (Mid-Atlantic -Fishery Management Council
             1990, 1991a,b), the status has shown some improvement with about a 70% and 50%
             increase in commercial and recreational landings, respectively, between 1990 and
             1992. However, the total reported commercial landings in 1992 (7,300 metric tons)
             were only about 40% of the total landings in 1979 when this fishery peaked (17,945
             metric tons) and 1992 recreational landings were only about 25% of the 1983 peak for
             this fishery (Terceiro 1993). In addition, the most recent National Marine Fisheries
             Service/Northeast Fisheries Science Center fishery-independent groundfish survey
             indices for spring are about one-fifth of those for the mid-to-late-1970s (Mid-Atlantic
             Fishery Management Council 1990, 1991a,b). Along with reduced abundance, the age
             structure of the population is severely compressed with relatively few individuals older
             than three years of age (Terceiro 1993). As a result, the spawning individuals are
             younger, presumably less fecund than older, larger individuals, and have fewer
             seasons to reproduce before being harvested.

             Despite the importance of summer flounder, there are significant gaps in our
             knowledge of the population dynamics of this species. For instance, several years of
             exceptionally poor recruitment throughout much of the range (Mid-Atlantic Fishery
             Management Council 1990, 1991a,b) suggest other factors besides fishing pressure
             affect summer flounder abundance, but these are poorly understood. Two major
             reasons why the patterns of recruitment are inadequately understood is that our
             knowledge of young-of-the-year habitats is poor, which may result in extremely
             variable young-of-the-year assessments. In addition, recent habitat-oriented studies
             have suggested that low temperatures encountered by juveniles in estuarine nurseries
             during the winter may impact both survival and subsequent year-class strength
             (Malloy and Targett 1991; SzedImayer et al. 1992). These kinds of results need to be
             summarized so that resource managers can effectively protect or enhance important
             nursery habitats. Thus, the purpose of this document is: 1) to synthesize the
             available literature (both primary and gray) on habitat parameters and habitat use
             patterns of all life history stages (egg, larval, juvenile, adult) of summer flounder








                2
                                                                                           Introduction


                throughout its range; 2) to incorporate unpublished data from federal, state and
                academic institutions into the synthesis; and 3) to provide a complete bibliography for.
                all aspects of summer flounder biology.

                SCOPE

                For purposes of this synthesis, we define habitat as where an animal lives (Odum
                1970). We interpret this broadly to include general distribution of all life history stages,
                as well as movements and selected aspects of the biology, such as those that appear
                related to habitat use or impacts on-habitats (anthropogenic effeqjs). We have
                cautiously interpreted prior generalizations, regarding, summer flounder habitat, and are
                particularly careful not toi,,e;@trapolate' from general statements concerning other closely
                related flQ..pnders (Paralichthvs.spp.). -Because'pertinent life history i-riformation is
                frequently lacking; we include-a brief.-sum.mary of our current understanding of the
                ,salient aspects. . 5iirice@ different. stqcks,or populations may have.-different habitat
                requirementsior patterns,of habitat use,we.also briefly, summarize what is known
                about. distribution and -stock @identification., Me, recogpizethat we still lack important.
                information, about the thabitat of-surnmer flounder.d.uring all life history stages.




























                Odum, E.P. 1971. Fundamentals of Ecology. W.B. Saunders Co., Philadelphia, PA










             Chapter 2
             Distribution and Life History                           Patterns


             GEOGRAPHICAL DISTRIBUTION


             Summer flounder occur in continental shelf and estuarine waters of the western North
             Atlantic (Fig. 2.1) from Nova Scotia, Canada (Bigelow and Schroeder 1953; Scott and
             Scott 1988), to Florida, United States (Ginsburg 1952; Gutherz 1967; Gilbert 1986;
             Grimes et al. 1989). They are most abundant from Cape Cod, Massachusetts, to
             North Carolina (Grosslein and Azarovitz 1982). In the north, commercial and
             recreational catches drop in the vicinity of Rhode Island', and occurrence north of Cape
             Cod Bay is described as extremely rare (Bigelow and Schroeder 1953). Distribution in
             the southernmost part of the range is not clear partly because summer flounder are
             not identified to species in the commercial fisheries, which treat them largely as by--@
             catch. Gilmore et al. (1981) list them as abundant in the Indian River Lagoon on the
             east coast of Florida, and Gilbert (1986) considers them to extend as far south as
             Sebastian Inlet, Florida.


             GENERAL LIFE HISTORY


             Summer flounder reproduce in the fall and perhaps into the winter. Detailed
             observations of the timing and location of spawning are available only for Georges
             Bank and the Middle Atlantic Bight (Morse 1981; Able et al. 1990; Fig. 2.1). There are
             no records of eggs from estuaries, but based on collections of planktonic eggs during
             the period from 1975 to 1985, spawning occurs during the seasonal offshore migration
             and is fairly equally distributed over the entire continental shelf (Fig. 2.2). Peak
             spawning occurs in October and November throughout the'Middle Atlantic Bight and
             Georges Bank, although December collections are lacking from some areas (Fig. 2.3).
             Spawning occurs over the continental shelf in the South Atlantic Bight as well,
             however data are largely lacking in the existing literature (Powles and Stender 1976).

             The planktonic larvae are reported over the Middle Atlantic Bight shelf but are not very
             abundant over Georges Bank (Fig. 2.4, 2.5). The earliest spawning and subsequent
             larval development occurs off eastern Long Island and on Georges Bank as early as
             September. By October, the larvae are primarily found on the inner continental shelf









                                4
                                                                                                                                                                                                   Distribution




                                                                                                                                                                                                    Cape
                                                                                     NOVA
                                                                                    SCOTIA


                                                                 INE
                                                                                                                                                                               ... @Zt

                                                                                            6e
                                                            EW HAMPSHIRE                                                   Cod Bay
                                                                      MASSA-
                                                                                            4C@                                                                                             . . ........
                                                                        HUSETTS
                                                                                                                                                                                       X:   @Co

                                                                                                                                                                                    ...... .. ......
                                                                RHODE                                                   ong Island
                                                                                            6el                                                                                             ......
                                                                     ISLAND                                                     Sound
                                                                                                                                                                               ............
                                                                                                                                                                              .............
                                                                                                                                                                                ..........
                                                                  NNECTICUT
                                                                                                                                                                                        Long Island
                                                          NEW YORK
                                                                                                                    nasquan River
                                                                                                                                                   th
                                                          NEW JERSEY                        7CP,                        at Say - Little            ira
                                                                                                                       Egg Harbor
                                                            LAWARE                                                 elaware Bay
                                                            MARYLAND                        3(f'
                                                                                                                                                                                       Cape
                                                                                                                           sapeake                                                      Charles
                                                            VIRGINIA                        72@                            Bay                                                .......... .
                                                                                                                                                                                            Cape
                                                                                                                               lbemarle                                                     Henry
                                                                                            3e                                     Sound

                                                                                                                               Pamlico                                            .@F:-X.%     Hatteras
                                                                                                                                  Sound
                                                                                                                                                                                            Cape
                                                                 NORTH                      7,01                                                                                               Lookout
                                                                    CAROLINA
                                                                                            3_                                                                                                 Cape
                                                                                                                                                                                                  Fear


                                                            SOUTH
                                                                CAROLINA                                          Charleston                                             ........
                                                                                                                           Harbor
                                                                                                                                                                             Y
                                                                              N             3
                                                                                            7                                                      VIC.

                                                      GEORGIA


                                                                             0 100
                                                                                                                       Indian River
                                                                                                                           Lagoon
                                                                                            2e'
                                                                          FLORIDA
                                                                                                                                             der;
                                                                                            78P:                                             let

                                                          UNITED                                                                                                  OSUMMER
                                                          STATES                            2e           Gulf of                                                  FLOUNDER
                                                          CANADA                            8d                  MeXICO                                            RANGE

                                                Figure 2.1.                East coast               of    the United States and Canada with features
                                                mentioned in the tex







                 Distribution
                                                                                                                             5



                       .44         f6           74            7'2           7'0
                                                                      ME

                                               e%
                                  EGGo
                                                                  NH-

                                                                              ................
                        42                                       MA                                            1`4 2A
                                            NY         CT        R I


                         40               NJ                                                                    40,

                                 DE

                                                  V                          IOM2       /rectangle,
                                                                                                                3.8,
                                                                                         0
                       V A                                                              .01-.25
                                                                                        .26-.50
                                                                                          51-1.0
                           N                                                             > 1.0

                                                             7-2             7.0            6.8
                                             7.4

                    Figure 2.2. Cumulative distribution and mean abundance of summer flounder eggs from National Marine
                    Fisheries Service (NMFS), Marine Resources Monitoring, Assessment and Prediction (MARMI         'AP)
                    offshore surveys during 1977-1984 (Sherman 1980 1986). Monthly to bimonthly plankton samples were
                    collected from Cape Sable to Cape Hatteras (@ig. 2.1) using 61 cm bongd'frames (Sibunka and
                    Silverman 1984; Morse et al. 1987). The 200 m and 1000 m contours are shown. (Figure from Able et
                    al. 1990).




                                                                                                                4 j










                                                                                                  Distribution




                           loo.                                         GEORGESBANK

                             0-:


                           100


                        W
                        L< loo                                    SOUTHERN NEW ENGLAND

                        Cn   0


                           loo
                        LL
                        0
                       04
                        E  100                                            NEW JERSEY

                             0-
                        (n


                           100-

                        U.
                        0
                        %  loo.                                     DELMARVA PENNINSULA

                        z
                             0-                             NS

                        z
                        W  100



                           100                                          VIRGINIA CAPES
                                                                      TO CAPE HATTERAS
                             0-:                            NS


                           100

                                JUL AUG SEP OCT NOV.DiC J@N           FiB MAR APR MAY JUN
                       Figure 2.3. Monthly abundance of summer flounder"eggs by region from National
                       Marine Fisheries Service (NMFS), Marine Resources Monitoring, Assessment and
                       Prediction (MARMAP), offshore surveys during 1979-81, 1984, and 1985. Monthly
                       to bimonthly plankton samples were collected from Cape Sable to Cape
                       Hatteras (Fig. 2.1) using 61 cm bongo frames (Sherman 1980, 1986; Sibunka and
                       Silverman 1984; Morse et al. 1987). Southern New England is the offshore region
                       between southeastern Cape Cod and northern coastal New Jersey. Delmarva is
                       the peninsula between Delaware and Chesapeake bays that is part of Delaware,
                       Maryland and Virginia. NS  no samples.







                               Distribution
                                                                                                                                                                                                                          7





                                                                                                                Z"
                                           ill             i4l             i2        i                           JER               .44  iG              i4          i2                iQ                               44@
                                   -44
                                         Paralichthys dontatus                                                                         Paralichthys dentatus
                                                  1977
                                                           -1987                                                                               1977-19B7
                                                Larvae     10 ma                                                                             Larvae 1,10 mz
                                                           0
                                                 EE
                                                                                                                                                        t

                                   42
                                                                                                                         42-       -42                                                                                 42-






                                                                                                                                                                                                                       40-
                                   40                                                                                    40-       40

                                                                                N                                                                                           N



                                                                                                                                                                        kilometers
                                                                                ters
                                                                                MSNQ
                                                                                  100                                                                                   0        100
                                                                           0
                                   38                                                                                    38        -3B                                                                                 38
                                                                           September
                                                                                                                                                                         October





                                                                                                                         36        5                                                                                   36


                                                           ?4          72                             BB
                                                                                                                       66-                              74         @2                 ?.o           B@B

                                           76              74              72        @0             68          //616                   76              74          7 2               70         68'           66
                                   -44                                                                                   44m       -44                                                     4,                          44-
                                          Paralichthys     dentatus                                                                     ParAchthys      dentatus
                                                                                                                                                        -1987
                                                 1977-1987                                                                                      1977
                                               Larvae / 10 mz                                                                                 Lar"e     / 10 Ma
                                                           10to                                                                                         0
                                                                                                                                                        A-10
                                                           11-100                                                                                       11-100
                                   42                                                                                    42        42                                                                ------            42






                                   40                                                                                    40        40                                                                                  40

                                                                               N                                                                                            N

                                                                                                                                                            X

                                                                           kJoraeLers                                                                                   kilometers
                                                                           0      10                                                                                    0        LOO
                                                                                                                         38        38
                                                                                                                                                        @:'                                                            3B
                                   -38
                                                                           November                                                                                      Oecember




                                              j
                                   36                                                                                    36                                                                                            35-


                                      7                    @4          @2             @0              68               66          ?-76                 74         @2                 7.0           6.8

                                   Figure 2.4. Monthly distribution and'm;ean abundance of summer flounder larvae from                                                                        National Marine
                                   Fisheries Service (NMFS), Marine. Resources Monitoring, Assessment and Prediction (MARMAP)
                                   offshore surveys during 1977-1987. Monthly to bimonthly plankton samples ,were collected from Cape
                                   Sable to Cape Hatteras (Fig. 2.1) using 61 cm                                     bongo frames (Sibunka and-S,ilverma.n 1984; Morse et al.
                                   1987). The 200 m contour is shown. (Figure by                                         M. P.     Fahay,       National        Marine Fisheries Se.rvice, Sandy
                                   Hook, New Jersey).









                                                                                                                                                                                                    Distribution










                                                                           i2            7,0           il'i                                 7-6             7-4           7-2           7-a         IS8
                                      .44     @6              7'4                                                  1'6       44      -44                                                      @4 1.
                                             Paralichthys dentatus                                                                        Parallchthys dentatus
                                                    1977-1987                                                                                     1977-1987
                                                   Larvae / 10 mz                                                                                 Larvae    10 mz
                                                              0                                                                                             0
                                                              1 10                                                           4?@                            1-10
                                                              11:100                                                                              @i        11-100
                                      42                                                                                     42      42                                                                                    42





                                                                            -6-
                                                                                                                             40      40                                                                                    40
                                      40

                                                                                  N                                                                                            N


                                                                                     ters                                                                                   kiloi ters
                                                                                                                                                                            0      1
                                      -38                                                                                    30      -38  P                                                                                38
                                                                               January                                                                                      February





                                      -313                                                                                                                                                                                 36.
                                                                                                                             35      -36


                                                          74                                                                                           @4
                                                                                          @O                                                                              @2

                                                                                         50            68-          169                                     i4            i2            7i10
                                              76              ?4           i2                                                                                                                                              44-
                                      44                                                                                             -44                                                       -11
                                             Paralichthys dentatus                                                                        Paralichthys dentatus
                                                                                                                                                  1977-1987
                                                     1977-1987
                                                   Larvae     10 m2                                                                               Larvae    10 m2
                                                              0                                                                                             0
                                                    Ed                                                                                            Ej        .1-10

                                      42                                                                                     42      42                                                                                    42






                                      40                                                                                     40      40                                                                                    40

                                                                                  N                                                                                            N


                                                                               kilometers                                                                                         ters
                                                                              a        100                                           -                                              Lao
                                      -36                                                                                                                                                                                  36
                                                                                                                             38      -38  a
                                                                                March                                                                                         April





                                      36                                                                                     36                                                                                            3B


                                                          74              @2                                               66          78              74                 72            70              68

                                      Figure 2.4 (continued).






                Distribution






                                                                          GEORGESBANK
                             100.


                              0-


                             100.

                        LU
                                                                    SOUTHERN NEW ENGLAND
                        U<.
                        cc   100'
                        M
                        Cn    0-

                        Cn
                        LL   100'
                        0
                       N
                        E                                                  NEW JERSEY
                             100-


                        W     0


                             100.


                        LL
                        0                                            DELMARVA PENNINSULA
                        M    100
                        W
                        M
                              0-:
                        z
                        z
                        <    100-
                        Uj


                                                                            VIRGINIA CAPES
                             100.                                         TO CAPE HATTERAS

                              0'                             NS


                             100

                                 JUL A6G SEP OCT NOV DEC JAN FEB MA'R A@R MA'Y JUN

                        Figure 2.5. Monthly abundance of summer flounder larvae by region from National
                        Marine Fisheries Service (NMFS), Marine Resources Monitoring, Assessment and
                        Prediction (MARMAP), offshore surveys during 1979-81, 1984, and 1985. Monthly
                        to bimonthly plankton samples were collected from Cape Sable to Cape
                        Hatteras (Fig. 2.1) using 61 cm bongo frames (Sherman 1980, 1986; Sibunka and
                        Silverman 1984; Morse et al. 1987). Southern, New England is the offshore region
                        between southeastern Cape Cod and northern coastal New Jersey. Delmarva is
                        the peninsula between Delaware and Chesapeake bays that is part of Delaware,
                        Maryland and Virginia. NS  no samples.








               to
                                                                                                Distribution



               betwe 'en Chesapeake Bay and Georges Bank. During November and December,
               when the larvae are most abundant in ever region, they are fairly evenly distributed
                                 i                         ' y
               over both the inner and outer portions of the shelf. By January'and February, the
               remaining larvae are primarily found on the middle and outer portions of the shelf. In
               April, the re.mai,ning larvae,,,are concentrated off, North Carolina.
               As the larvae  undergo   eye migration during transformati  .on (Fig. 2.6), they enter
               estuarine nu rsery areas (Bu rke et al.. 199 1;, Keefe and Able f 993). Based on data f rom
               the Great Bay-Little Egg Harbor estuanne,system in southern New Jersey (Fig. 2.7a),
               this occurs just as the right eye is migrating across the top of the head (transformation
               stage G th 'rough H+) at sizes of 8 - 16 mm SL [standard length] (Fig. 2.8). They then
               leave the water@ column and settle to the bottom where they begin to bury in the
               substrate and complete development to the juvenile stage (Keefe and Able 1994). The
               morphological transition from larva to juvenile lasts beyond initial settlement, making
               the distinction between lato;'Iarva and early juvenile somewhat arbitrary. We suggest
               that .summer flounder be considered juveniles,when they have reach the I stage (Fig.
               2.6), when the pectoral fin resembles that of adults, and when the scales are formed.
               Further, we suggest that those individuals that have not completed ijye migration,
               pectoral fin and scale formation be'. referred tbas transforming larvae, as Burke (1991)
               has done. If larvae enter the estuary When water temperatures are low, mortality can
               occur (SzedIma   yer et al. 1992) or transformation and growth can be delayed for weeks
               (Keefe and Able 1994) until warmer temperatures allow continued. growth and
               development. The ability to delay transformation may help explain the seeming
               disparity between a peak of,,,spawning in October and November and the occurrence
               of transforming larvae in estuaries over a protracted period from -October through April.

               Recently, settled juveniles can be found, in a variety of habitats, but.by summer many
               are found in salt Marsh creeks (Szedlmayer'et 'al. 1992; Wenner etal. 1990; Burke et
               al. 1991; Rountree and Able 1992a,b; Rountree and Able 1993). Growth during the
               summer'is exceptionally fast, and at least in some estuaries can average 1.9 mm per
               day (Szedltaye.r et al. 1992). By September or October, the individuals that were
               spawned the previous fall have reached 230 to 300 mm TL [total length] (Able et al.
               1990; SzedImayer et al. 1992; Fig. 2.9). These fast growth rates appear to be common
               to most estuarine nurseries (Almeida.et al. 1992). At this time and size, many of these
               juveniles begin to move out of.the estuafries and onto the continental shelf (Able et al.
               1990; SzedImayer and Able 1993). The adults -follow a similar pattern (Fig. 2.10, 2.11),
               migrating in the fall to spawn over the continental shelf. They overwinter near the
               edge of the shelf -and then migrate back into inner continental shelf and estuarine
               areas the following spring.










                                                                                                                                                             0
                                                                                                                                                             U)


                                                                                                                                                             Cr


                                                                                                                                                             0






                         Pre-transformation                        F -                           F






                              . . . . . . . . . . .







                                     H -                           H                            H +

                       Figure 2.6.  Classification of transformation    stages  for summer flounder based on degree of          eye migration
                       (adapted from Keefe and Able 1993). The right and left eyes are bilateral and symmetrical in pre-transformation
                       individuals. At the first.stage of transformation, F -, the eyes are bilateral but asymmetrical with the right eye just
                       dorsal to the left eye. By Stage G, the right eye is visible from the left side of the fish. Stage H - differs from G in
                       that the cornea of the eye is visible from the left side of the fish. At Stage H, the right eye has reached the dorsal
                       midline. By Stage H +, the right eye has reached the left side of the head but has not yet reached its final resting
                       place. At Stage I , the eye is set in the socket and the dorsal canal has closed.








                                            12
                                                                                                                                                                                                                                                                         Distribution




                                                                                                         .........................................                                           .... ....
                                                                                                                                                                                             .......  ::...:                                                     ............................
                                                                                EAST:                                                                                                                                                       .. ........
                                                                                                         .... ..........
                                                                                                                                            ........ . . .
                                                                                                                                            ......                                                                          .. ................
                                                                                COAST                                                       .......... ......     : I A&                                                    .
                                                                                                         ............
                                                                                UNITED:
                                                                                                                                                                                                                                                                 ............ ......
                                                                                                         ..........                                                                                                         ... ...                              .. ..... . ..
                                                                                STATES                                                                                                                                                                           . .. .... ... .
                                                                                                                                                                                                                                                                                BLOCK
                                                                                                                                            ........ ..
                                                                                                                                                                                                                                                                                      ISLAND
                                                                                                                                            ARSHELDEA
                                                                                                                                            M                                                                                    LONG ISLAND
                                                                                                                                                                                                                                                                                          SOUND
                                                                                NEW
                                                                                                                                                                                                                                                                         %
                                                                                                                                                                                                                                 SOUND
                                                                                                                                                                  ISLAND
                                                                                JERSEY

                                                                                                                                                                                                                                                                         COAST..
                                                                                                                                            E
                                                                                                         ... ...                            gg:,
                                                                                .......................... ..... ...
                                                                                . .......... ...                                                                                                       .....
                                                                                ......                                                      Harbor                                                                                                                       STATW
                                                                                                         ............
                                                                                                                                                                                                                                                                 ............
                                                                                                                                                                                                                                                                 'Or
                                                                                                                                                                                                                            W    bAK

                                                                                :X:
                                                                                                                                                                                 ::,X
                                                                                .......... . .. ... ..
                                                                                X.-                                                         pshead
                                                                                ...............
                                                                                                                                            .@i@@i"Creek
                                                                                .......                                                                                                                                                                                               NEWYC A
                                                                                . .......                B                                                                                                                                                                            CONNEC
                                                                                                         ay
                                                                                                         @@-":Q,.,RUMFS
                                                                                ..............
                                                                                ... ..........
                                                                                                                                                                                                                            ...... ... .. .. ....
                                                                                                                                                                                 EAST:
                                                                                                                                            Little
                                                                                .. chooner

                                                                                .. . ... ....                                                                                                                               ......
                                                                                                                                            Egg                                  TEV:                                       Albernarle Sound
                                                                                                         ........... .. ...
                                                                                                         Creek
                                                                                                                                                                                 UM
                                                                                                         .... .. ...... ..
                                                                                                                                                                                 st, TES.,:@
                                                                                                                                                                                                                                                                 X..
                                                                                ..........               .... .........                     Inlet
                                                                                                                                                                                                                                                                                      Roan
                                                                                                                                                                                                                            ...... ......                                                     oke
                                                                                ............
                                                                                ...............          ....
                                                                                                                                                                                                                                 X                               zi*i
                                                                                                                                                                                                                                                                 .......              island
                                                                                                                                                                                                                                                                 ..............
                                                                                                                                                                                                                                 .................. .
                                                                                                                                                                                                                            x, ......    ...............
                                                                                ............. ...        ........... ....
                                                                                                                                                                                                                            . ..... .. .. ... ---
                                                                                ......                   ...... .
                                                                                ............             .... ...
                                                                                                         .. . ...... ...
                                                                                                                                                                                                                                                                 ............ ..      regon
                                                                                                         .. .....                                                                                                           ........................
                                                                                                                                                                                                                            ...                                  . . . ...... .
                                                                                                                                                                                                                                                                 ......%              .    Inlet
                                                                                                                                                                                                 NORTH                                                           . .. ....... .
                                                                                .#Y
                                                                                                                                                                                          OCAROLINAf@@.
                                                                                                                                                                                                                                                                 ..........
                                                                                                         ..... ...........
                                                                                                                                                                                                                                         X.                      ..................

                                                                                SIM
                                                                                an.                                                                               NEW:
                                                                                                                                            P                                                                                                                    ..........
                                                                                                                                                                                                       VW
                                                                                                                                                                                                      ...................................
                                                                                                                                                                                 . .......... :..
                                                                                                                                                                                 . ............ .....
                                                                                ......... .
                                                                                ................                                                                                                                                                                                           Outer
                                                                                                                                                                                 X
                                                                                . .. .......
                                                                                                                                                                                                                                                                 ........ Pamiko
                                                                                                                                                                                             X":
                                                                                                                                                                                                                                                                                            Banks
                                                                                .. ...... . .Absecon Inlet                                                                                                                                                       Sound
                                                                                                                                                                  Study                                                                                               East
                                                                                ATLANTIC CrrY                                                                                                                                            ......
                                                                                                                                                                                 WW"                                        . .... .
                                                                                                                                                                                                                                                                                      -::@Cape
                                                                                                         ...............................    :X
                                                                                :EAST.:.                                                                                                                                                    Bluff Shoal
                                                                                                                                                                                                                            Parntico
                                                                                                         ........                                                 JU                                                                                                                  Hatteras
                                                                                COAST                                                                                               .......                                 Sound
                                                                                UNITM
                                                                                                               .... ...........
                                                                                ,STATES::
                                                                                                                                            X:: -:-..-.X:         :X ...........
                                                                                                         .............                                                                                                           West
                                                                                                                                                                                                                                                                 x
                                                                                                         .......                            ....................... %.-....,                                                                                     -`Ocracoke Inlet
                                                                                                         ..........
                                                                                                                                                                                                                                                                                           N
                                                                                                                                            ...... ........ .. ..
                                                                                                                                            ... . .. ...... ..
                                                                                                               . .......                    ............ .................
                                                                                                               : :-                         . 1: ....%%............... ...
                                                                                                         ........ . .. .....                                      ... . ..
                                                                                                               . ...... .                                         ...%. .......
                                                                                                               .......... ..                                      ............
                                                                                                               ..........
                                                                                                                                                                                                                                         ..."Banks
                                                                                                         ......... ..                                             .........      . ...
                                                                                                                                                                  ........ ..                                               ... ..........
                                                                                SOUTH                                                                                                                                                    D                       i Inlet
                                                                                                         .. ..........
                                                                                                                                                                  ;:;:N .....
                                                                                                                                                                                                                                         rum
                                                                                C
                                                                                AROLINA
                                                                                                                                                                  X.
                                                                                                         X
                                                                                                                                                                  X.X.
                                                                                .................. .                                                                                    .............                       .... .......
                                                                                .... . ..... ..... .
                                                                                ..................       X.
                                                                                                                                                                                                                            .70-core Sound
                                                                                                                                                                  ............
                                                                                                                                                                  .............
                                                                                                                                            ......... ....
                                                                                                                                                                  x,
                                                                                .................
                                                                                ........                                                    ... ....                                                                                     S aund
                                                                                .......... ....
                                                                                ..............
                                                                                                                                                                  . ............        Beaufort
                                                                                                                                                                                                                                                                                      0 10 20
                                                                                                                                                                  X,X.:.:.X:                                                Cape
                                                                                :,'Ah
                                                                                                                                                                                         Inlet B.
                                                                                                                                                                                 90gue                   rden's
                                                                                .. ...........                                              ....... ... .
                                                                                                                                                                                                                            Lookout
                                                                                                                                                                                                                                                                                      k1lorneters
                                                                                ..........                                                                        . ............ Sound               Inlet

                                                                                ..........
                                                                                                         ............
                                                                                                                                                                                 Figure 2.7. A.                             Great        Bay                     -Little Egg          Harbor
                                                                                                                                            .. .. ......... .
                                                                                                         .... ......
                                                                                ... ..                   ...... ..........
                                                                                                                                                                                 estuarine            system                in southern                          New Jersey with
                                                                                                                                            X.
                                                                                                         . ..... ..... ..........           .....                 ....           h
                                                                                                         ......                                                                  t e location of                            Rutgers University Marine Field
                                                                                ......                                                                                           Station           (RUMFS) indicated.                                                 Study sites
                                                                                                         ......                             ........ .....                       referred        to in
                                                                                                                                                                                                                            the text are indicated by a circle:
                                                                                ........ ........ ...................
                                                                                                                                            ...........
                                                                                ........................                                    ......
                                                                                                         ..............
                                                                                                               .......                                                           B. Charleston                              Harbor estuarine system in South
                                                                                                         ..............
                                                                                .......                  ........... .
                                                                                                                                                                                 Carolina. Study creeks mentioned in the text are
                                                                                ..............
                                                                                                                                                                  ... . ........ .
                                                                                ............             . ..........
                                                                                                                                                                  ...            indicated by circled numbers and are as follows;
                                                                                                         ......... ...
                                                                                                                                                                                 1) Pita Creek,                             2) Lachicotte Creek, 3) Beresford
                                                                                ........                                                                                         Creek, and 4)                              Inlet Creek: C. Long Island
                                                                                ...........
                                                                                Charleston                                                                                       Sound system                               bordered by New York and
                                                                                                         Harbor@                                                                 Connecticut:                               D. Pamlico-Albemarle sounds
                                                                                                                                                                                 estuarine system in North Carolina.
                                                                                                                                                                                                                            N






                     Distribution
                                                                                                                                                               13




                                                 A.    80-
                                                                                                 ............
                                                                           LITTLE
                                                                                                 ......... .
                                                                                                 ............
                                                                   SHEEPSHEAD
                                                                                                 .. ..........
                                                       60-
                                                                           CREEK
                                                 Cn
                                                                                                       .. . ..........
                                                                  Nov 1990 - Nov 1991:
                                                                                                           ............
                                                                                                     .... ..........
                                                                                                 ........... .. .........
                                                                                                       . .........
                                                       40-
                                                                           n    400

                                                                                                                   ..... .....
                                                         0-
                                                                                        ..........


                                                 >     2
                                                 z                            F77771
                                                 LL    80-
                                                 0         -      SCHOONER CREEK
                                                 M                Feb 1990 - Apr 1992
                                                 LU    60-
                                                                           n     77

                                                       40-
                                                 z
                                                       20-
                                                        0 -F=                 FM                 Mill      ...... I
                                                            8-8.9            10-10.9             12-12.9          14-14.9
                                                                           STANDARD LENGTH (mm)


                                                 B.       100-
                                                                         LITTLE
                                                                                            ............
                                                                  SHEEPSHEAD
                                                           80-
                                                                         CREEK
                                                           60-    Nov 1990 - Nov
                                                                                            ............
                                                                                            .. ........
                                                                           1991
                                                           40-           n    400

                                                           20-
                                                    z
                                                                                                          ..............
                                                    W             EM
                                                              0--
                                                          100-
                                                                  SCHOONER CREEK
                                                           80-    Feb 1990 - Apr 1992
                                                    LL                             51
                                                           60-

                                                           40-

                                                           20-

                                                              0-
                                                                    G             H-           H          H+            I
                                                                           TRANSFORMATION STAGE


                                  Figure 2.8. A. Length-frequency, and, B.. stage-frequency distributions for
                                  transforming summer flounder larvae collected durin Rutgers University
                                  Marine Field Station ichthyoplankton surveys in two Uat Bay-Little Egg
                                  Harbor estuarine system marsh creeks (Fig. 2.7A). Sampling was conducted
                                  with 1.0 m and 0.5 m plankton nets with I mm mesh. See Fig. 2.6 for
                                  transformation stage descriptions.








            14
                                                                           Distribution



                    30--
                                        NOV-APR                       21 - 26 JUL
                     .20-:                n  65                          n  21
                    10-
                     01
                    30--
                                        187 27 MAY                    2 - 10 AUG
                                          n  7                          n  106
                    20":

                    10-

                     0-
                >'  30--
                0                       14 - 15 JUN                   17 - 25 AUG
                z                        n  114                         n = 125
                    20,_:

                    10__
                w

                LL   0
                    30--                28 - 30 JUN               30 AUG - 08 SEP
                                          n  38                       n = 92
                    20-

                    10
                     01      X,
                    30--                11 - 14 JUL                   17 - 22 SEP
                                          n = 26                         n  50
                    20--

                    10-
                     oa                                             JA jj
                                                                     I I I   I r-1
                        0   100    200   300 400      0    100   200 300 400

                                           TOTAL LENGTH (mm)
                      Figure 2.9. Length-frequency distributions for young-of-the-year summer
                      flounder collected during 1988-1989 sampling in the Great Bay-Little Egg
                      Harbor estuarine system (Fig. 2.7A; adapted from Szedlmayer et al. 1992).






               Distribution
                                                                                                                           15





                                                                                 r__4
                                             ...........


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

                                              Q


                                           .. .......






                                                                            MEAN NUMBER OF
                                                                              INDIVIDUALS/
                                                                          ONE-HALF HOUR TOW
                                                                                  0 to 5
                                                                                 6 to 10
                                                                                 11 to 25
                                                                                26 to 50
                                                                                   50 +


                                                                            SPRING





                                                                                     X













                                                                            AUTUMNL


                               Fig. 2.10. Seasonal   distribution and abundance of summer flounder from
                               National Marine Fisheries Service (NMFS), Northeast Fisheries Science
                               Center (NEFSC) grdundfish survey& for spring (1982-1960) 'and a u-tumn
                               (1981-1990). Sampling was conducted by stratified random design in the
                               Gulf of Maine to Nova Scotia, on Georges Bank, and in the Middle Atlantic
                               Bight (Fig. 2.1) using 0.5 hour tows of a #36 Yankee trawl With a 12.7 mm
                               mesh liner in the codend . The 200 m and 1000 m contours are shown.
                               (Adapted from maps provided by T. Azarovitz and B. O'Gorman, National
                               Marine Fisheries Service, Woods Hole Laboratory, Massachusetts).








                   16
                                                                                                                Distribution



                                                                                                     A?


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




                                                  ?








                                                         AUTUMN                                 SPRING
                                                        MIGRATION                             MIGRATION












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

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

                                    3 ii: i ix@ X -,*

                                            . . . . . . . . . . . .:.::.:






                                    .. ........

















                                         . . ..........
                                         ............ . . . . . . . . .
                                    =
                                            2.11. Synopsis  of results  of tagging studies on   adult summer
                                            r depicting, A. offshore mov'ement from summer feeding grounds
                                    and, B. inshore movement from overwintering grounds. *indicates
                                    exact overwinter location was unknown; i.e., fish were tagged in an
                                    estuary in spring and were recaptured in an estuary the following spring. ?
                                    indicates no data available. Data are drawn from the following sources:
                                    Westman and Neville 1946); Hamer and Lux (1962); Poole (1962);
                                    Murawski (1970a); ScJetl (1984); Mercer et al. (1987); Desfosse et al.
                                    (1990); Jesien and Hocutt (1991).







              Distribution
                                                                                                     17



              STOCK IDENTIFICATION

              There have been, and continue to be, a number of attempts to resolve the identity of
              summer flounder stocks with a variety of techniques. To date, there have been three
              basic interpretations. First, the results of some studies have suggested that there are
              two major stocks, one in the Middle Atlantic Bight and another in the South Atlantic
              Bight. Many of these studies, beginning with Ginsburg (1952), have used meristic and
              morphometric techniques (Smith and Daiber 1977; Wilk et al. 1980; Fogarty 1981).
              However, other investigators (R. Jesien and C. Hocutt, pers. comm.) have been
              concerned with allometric effects on the analyses by both Wilk et al. (1980) and
              Fogarty (1981) because northern and southern study populations were of different
              sizes.


              A second interpretation identifies two Middle Atlantic Bight-based stocks. One stock
              appears to make a consistent offshore migration in the winter and return to estuaries
              and inner continental shelf waters in the summer. The second population appears to
              spend the summer in estuaries and inner continental shelf areas from Virginia to
              Maryland but overwinters near Cape Hatteras. This latter population has been
              referred to by some as the trans-Hatteras stock. This inte  rpretation can be derived
              from electrophoretic (Van Housen 1984), meristic and morphometric (Delaney 1986)
              analyses and tagging studies (Holland 1991). Both of these potential stocks occur in
              Chesapeake Bay and along the eastern shore of Virginia and perhaps Maryland in the
              summer (J. Desfosse, J. Musick, R. Jesien, C. Hocutt, pers. comm.). A third
              interpretation is a combination of the previous two in that it recognizes three stocks,
              one from the Middle Atlantic Bight, one from the South.Atlantic Bight, and the trans-
              Hatteras stock. Evidence to support the distinction in local populations north and
              south of Cape Hatteras is based on the tagging studies by the North Carolina Division
              of Marine Fisheries (Mercer et al. 1987; Monaghan 1992).

              The resolution of these different interpretations will probablynot be immediate, but
              because of the poor status of summer flounder populations (Mid-Atlantic Fishery
              Management Council 1990, 1991a,b), there is considerable focus on this problem.
              Current emphasis is on tagging studies in Maryland, Virginia and North Carolina.
              Whether or not continued study eventually resolves stock identity, two results from the
              tagging studies are of interest. First, studies in which fish have been tagged in
              estuarine areas show that there are a large number of returns to the same estuary the
              subsequent summer (Westman and Neville 1946; Poole 1962; Hamer and Lux 1962;
              Murawski 1970a; Desfosse et al. 1990; Holland 1991; Jesien and Hocutt 1991; Jesien et
              al. 1992; Monaghan 1992). Most of these studies are from the Middle Atlantic Bight
              where these fish presumably overwinter along the edge of the continental shelf or
              closer to shore in the south. Second, there is a trend for some fish to be found to the
              north and east of the original capture sites.










             Chapter 3
             Eggs


             INTRODUCTION


             Summer flounder eggs are buoyant (Smith 1973a,b), and ascend from the benthic
             spawning habitat on the continental shelf to upper ocean waters. Their distribution is
             a useful means of defining general limits of spawning.

             SPATIAL AND TEMPORAL DISTRIBUTION

             Summer flounder eggs have been collected over adult spawning grounds from inner to
             outer continental shelf waters as early as September in northern regions (Smith
             1973a,b; Able et al. 1990) and as late as January (Able et al. 1990) and February
             (Smith 1973a,b) in southern regions (Figure 2.2, 2.3). From Georges Bank to Cape
             Lookout, peak concentrations occur primarily in October and November (Able et al.
             1990), however Smith (1973a,b) reported a December peak off Cape Hatteras in 1965.
             In this region, stratified sampling to 33 m depth collected most eggs in the upper 15 m
             of the water column (Smith 1973a,b). Information on egg distribution south of Cape
             Lookout is not available.


             TEMPERATURE

             Vertical movement of the eggs from the bottom after spawning may expose them to as
             much as a 150C change in water temperature, but the available data suggest that they
             can tolerate such a range. Between Cape Cod and Cape Lookout, eggs were
             collected in temperatures from 9.1 - 22.90C, with most occurring in water temperatures
             of 13 - 17.90C (Smith 1973a,b). Laboratory studies have shown that egg incubation
             time and water temperature are positively correlated between 5 - 210C, and that in this
             range, normal development and hatching occurred from two to nine days (Smith and
             Fahay 1970; Smigielski 1975; Johns and Howell 1980; Johns et al. 1981). Further
             information is available from a study on the potential effects of entrainment in once-
             through cooling systems of power plants (Smith et al. 1979). Late embryo stage eggs
             were exposed to sudden increases in temperature (pre-experiment acclimation
             160C; experimental shock exposures = 23, 26, 29, 32, and 350C) for time periods
             ranging from 3 - 180 minutes at each shock level, and survivorship was high. One
             hundred percent mortality occurred only at 350C for exposure times at and above 90
             minutes.











            Chapter 4
            Offshore Larvae


            INTRODUCTION


            After hatching over the continental shelf, larvae are planktonic, symmetrical in form,
            and dependent on their yolk sacs for nutrition. Under usual temperature and salinity
            conditions, the yolk sac is fully absorbed within four days and feeding begins
            (Smigielski 1975; Johns et al. 1981). Larval development in the Middle Atlantic Bight
            occurs during the autumn breakdown of the thermocline and the resultant plankton
            bloom (Morse 1981). Transformation (Fig. 2.6) begins offshore and may be
            accompanied by behavioral changes that aid movement into estuaries (Burke 1991)
            where permanent settlement occurs.

            SPATIAL AND TEMPORAL DISTRIBUTION

            Offshore larval distribution (Fig. 2.4) is similar to that for eggs (Fig. 2.2). Between
            Georges Bank and Cape Lookout (Fig. 2.1), larvae have been collected as early as
            September in the northern regions (off Cape Cod) and as late as May in the south (off
            North Carolina) (Smith 1973a,b; Smith et'al. 1975; Bolz et al. 1981; Able et al. 1990).
            Data on larval occurrences south of Cape Lookout are scarce, partly because
            identification is complicated by thepresence of the larvae of two and sometimes three
            other species of Paralichthys (Powles and Stender 1976).

            TEMPERATURE

            Larvae have been collected over a wide range of water temperatures (0 - 23.10C)
            between Cape Cod and Cape Lookout, with most occurring at temperatures of 9.0 -
            17.90C (Smith 1973a,b). In the laboratory, yolk sac larvae died at temperatures below
            I 10C, but those held above I 11C developed normally (Johns and Howell 1980; Johns
            et al. 1981). Studies on the potential effects of power plant cooling systems to
            summer flounder larvae provide further information. The ability of newly hatched yolk
            sac larvae to avoid predation was actually enhanced after a +100C thermal shock
            (base 15. 1 OC; Deacutis 1978), however mortality resulted from shock exposure to 320C
            (Hoss et al. 1974).









               22
                                                                                  Offshore Larvae



               SUBSTRATE

               Planktonic larvae may begin making excursions to the bottom before entering
               estuaries (K.W. Able, unpubl. data). Small numbers of transforming larvae have been
               collected with a small-mesh beam trawl during autumn 1991 from an inner continental
               shelf site (15 m depth) off New Jersey. These larvae were in early transformation
               (pre-transforming to stage H, 11.4 - 15.4 mm SL; Fig. 2.6) and had coloration typical of
               larvae that occurred in the plankton except some of the melanophores were stellate
               (expanded), which made them appear more pigmented than normal. We suspect that
               these larvae had settled on the substrate intermittently on their way into the adjacent,
               estuary (Fig. 2.8) because larger, later-stage summer flounder larvae were seldom
               found at this site, even though large numbers of other recently settled flatfishes
               (Etropus Scophthalmus were often collected (K.W. Able, unpubl. data). There is no
               evidence that permanent settlement occurs on the continental shelf (Able et al. 1990),
               although benthic sampling with the appropriate small-mesh gear has been infrequent.










            Chapter 5
            Estuarine Larvae


            INTRODUCTION


            Summer flounder larvae entering estuaries are undergoing morphological
            transformation and are probably changing physiologically as well in order to cope with
            fluctuations of salinity in the estuary. Movement into the estuary may involve
            intermittent settling to test the substrate and larvae may take advantage of tidal stream
            transport prior to permanent settlement in the estuary. Because of the lack of
            developmental (stage) information in much of the literature, here we refer to all pelagic
            individuals as larvae and to all benthic individuals that have not achieved sexual
            maturity as juveniles.

            SPATIAL AND TEMPORAL DISTRIBUTION


            Summer flounder larvae move from offshore to shallow estuarine habitats to complete
            transformation and permanently settle (Keefe and Able 1993). In Middle Atlantic Bight
            estuaries from Long Island Sound to as far south as Chesapeake Bay, movement of
            larvae into estuaries has been reported to occur from October through April (Merriman
            and Sclar 1952; Olney 1983; Olney and Boehlert 1988; Able et al. 1990; Szedimayer et
            al. 1992). The only published report of the larvae in an estuary in summer (Herman
            1963) is questionable (Smith 1973a,b; Able et al. 1990). In the Great Bay-Little Egg
            Harbor estuarine system (Fig. 2.7a), larvae have been sampled in two marsh creeks
            near Little Egg Inlet, Little Sheepshead and Schooner creeks, during a five-year
            period (Figs. 5.1, 5.2). At Little Sheepshead Creek, temporal occurrence of larvae was
            sporadic within and between years. In the winters of 1989-1990, 1990-1991 and 1992-
            1993, most larvae occurred in January through March while few, if any, occurred in the
            fall. However, in the winter of 1991-1992, they were clearly more abundant in October
            through January. In Schooner creek, larvae were most abundant during December
            through March, although fall sampling did not occur in some years (Fig. 5.2). There
            seems to be some very general correspondence in abundance between these sites
            with the peaks in February 1990, February 1991, and December 1991 - February 1992
            co-occurring at each creek. Earlier sampling (1962-1972) in nearby Manasquan Inlet
            collected all larvae during October through December (Able et al. 1990). In the South
            Atlantic Bight, movement into the estuary occurred from January through April with
            abundance peaks in February. and March in North Carolina and South Carolina
            (Weinstein 1979; Bozeman and Dean 1980; McGovern 1986; Hettler and Chester 1990;









                        24
                                                                                                                                   Estuarine Larvae



                                                             0.020                                          1989
                                                             0.016

                                                             0.012

                                                             0.008


                                                             0.004

                                                             0.000


                                                             0.020.                                      1989-1990
                                                             0.016@


                                                     co      0.012-
                                                      E      aow.

                                                             0.004

                                                             0.000


                                                             0.020.,
                                                                                                         1990-1991
                                                      Z      0.016-
                                                      LL     0.012-
                                                      0
                                                      M      0.008-
                                                      ILLI
                                                             0.004

                                                             0.000



                                                      Z      0.020.
                                                      -<                                                 1991-1992
                                                      UJI    0.016.

                                                             0.012-

                                                             0.008-

                                                             0.004

                                                             0.000


                                                             0.020-                                      1992-1993
                                                             0.016.


                                                             0.012-

                                                             0.008-
                                                             0.004  1-
                                                             0.000 -
                                                                    'OCT'NOV      DEC 'JAN 'FEB 'MAR 'APR'MAY'TUN

                                            Figure 5.1.      Catch per unit effort for summer flounder larvae moving into
                                            an estuary based on Rutgers University Marine Field Station
                                            ichthyoplanKton survey at. Little Sheepshead Creek (Fig. 2.7A) from
                                            February 1989 through May 1993. The sampling site is a broad creek
                                            that cuts through the salt marsh peninsula between Great Bay and Little
                                            Egg Harbor and recieves Atlantic Ocean water on flood tides. Two
                                            plankton nets (1 meter diameter hoop, 1 mm mesh) were deployed once
                                            a week during the night flood tide for 3 to 5 consecutive 0.5 hour sets.
                                            Volume of water strained was determined using flow meters. July,
                                            August and September were sampled but are omitted because no
                                            summer flounder were collected in these months in any year.






                     Estuarine Larvae
                                                                                                                                                       25






                                                          0.38'
                                                          0.36'                                  1990

                                                          0.08,

                                                          0.06


                                                          0.04-


                                                          0.02


                                                          0.00

                                                          0.12
                                                          0.10                            1990-1991
                                                     W
                                                     _j
                                                     CL   0.08


                                                          0.06


                                                          0.04
                                                     M
                                                     M
                                                     0.   0.02

                                                          0.00
                                                     W    0.12-
                                                     >                                    1991-1992
                                                     M    0,10.


                                                          0.08
                                                     0    0.06
                                                     M
                                                     LU
                                                          0.04

                                                     M    0.02
                                                          0.00*0
                                                          0.12- -
                                                          0.10.                            1992-1993

                                                          0.08.


                                                          0.06.


                                                          0.04


                                                          0.02
                                                          0,00LO *"An *0
                                                                                             112 S4511-2"3 4 5112 3'
                                                                 NOV DEC JAN FEB MAR APR MAY

                                       Figure 5.2. Catch per unit effort for summer flounder larvae moving into an
                                       estuary based on Rutgers University Marine Field Station ichthy6plankton survey
                                       at Schooner Creek (Fig. 2.7A) from late-February 1990 through mid-May 1993.
                                       The sampling site is a small (1 km) polyhaline marsh creek that terminates in
                                       high marsh. A stationary plankton net (0.5 m diameter hoop, 3.0 mm mesh) was
                                       fished in the lower main creek continuously (day and night) for 1 to 7, 24 hour
                                       periods each week. Data are summarized for 5 sample periods during each
                                       month where the first four periods represent 7 days and the fifth period includes
                                       the 2 or 3 days remaining in that month. Months not represented were not
                                       sampled except for several days in late October 1992 during which time no
                                       summer flounder were collected. Unconnected points indicate breaks in
                                       sampling.








               26
                                                                                     Estuarine Larvae



               McGovern and Wenner 1990; Warlen and Burke 1990; Burke et al. 1991). The
               consistency in transformation stages and sizes at which larvae enter Middle Atlantic
               Bight and South Atlantic Bight estuaries (Burke et al. 1991; Keefe and Able 1993)
               suggests that transforming larvae develop behaviors that first allow transport into the
               estuaries from offshore during early stages (Burke 1991) and then, during later stages,
               aid up-estuary movement to settlement habitats, perhaps by tidal stream transport as
               for Paralichthys spp. (Weinstein et al. 1980a). This is discussed further below (see
               Substrate and Behavior).

               TEMPERATURE AND SALINITY

               Temperature appears to be the most significant environmental factor controlling
               duration of transformation and thus, perhaps, timing of larval movement into estuaries.
               Late-development larvae have been collected in water temperatures ranging from 0 -
               130C in Middle Atlantic Bight estuaries. (Olney and Boehlert 1988; Wyanski 1990;
               Szedlmayer et al. 1992), from 8.4 - 23.40C in South Carolina (McGovern and Wenner
               1990) and from 2 - 221C in North Carolina (Williams and Deubler 1968a). Larvae
               collected in Little Sheepshead Creek (Figs. 2.7, 5.1) typically occurred at temperatures
               of 0 - 12.OOC. In Schooner Creek (Figs. 2.7, 5.2), larvae were collected at -2.0 - 140C.
               In the laboratory, however, water temperatures less than 20C killed transforming larvae
               (Szedlmayer et al. 1992). In another laboratory study of the same population, mortality
               of mid-transformation larvae held at 40C was significantly greater than those held at
               100C (Keefe and Able 1993). Thus, estuarine larvae suffer increased mortality based
               on both the severity and duration of cold water temperatures. Further, duration of
               transformation was dependent on ambient water temperatures, ranging from 25 days
               at average temperatures of ITIC to 93 days at average temperatures of 6.60C. Data
               for a South Carolina estuary suggest that a milder winter may mean earlier movement
               into the estuary (Cain and Dean 1976; Bozeman and Dean 1980).

               Larvae moving into estuaries are reported from a wide range, of salinities. In the two
               New Jersey marsh creeks (Figs. 5.1, 5.2), larvae occurred at salinities ranging from 20
               - 33 ppt. In North Carolina, larvae were collected throughout the estuary at salinities
               ranging from 0.02 - 35.0 ppt (Williams and Deubler 1968a). In a South Carolina study,
               larvae occurred from 0 - 24.7 ppt (McGovern and Wenner 1990).

               SUBSTRATE AND BEHAVIOR

               The estuarine movements of transforming larvae may be accomplished by vertical
               migrations between pelagic and benthic habitats (Weinstein et al. 1980a,b). In North
               Carolina, summer flounder and other species of Paralichthys responded to diel and
               tidal cycles by migrating to surface waters on night flood tides (Weinstein et al. 1980a).
               Many others have described increased catches of transforming larvae during night






            Estuarine Larvae
                                                                                                  27


            flood tides (Deubler 1958; Williams and Deubler 1968a,b; Olney and Boehlert 1988;
            Hettler and Chester 1990) and very high catches during new moon periods (Williams
            and Deubler 1968.a,b; Hettler and Chester 1990). Weinstein et al. (1980a,b) suggested
            that this behavior allows larvae to enter shallow habitats in marsh creeks and shoals
            where they are retained by an ebb-tide response of seeking the substrate. More
            recent behavioral data support these observations and indicate that transforming
            larvae may test the substrate before permanent settlement. Larvae observed in.the
            laboratory, in the absence of tidal currents, swam in the water column more often at
            night (Keefe and Able 1994). These larvae swam with their bodies held vertically, sank
            with their bodies held horizontally, and were capable of periodically resting on the
            substrate on their right sides even before completing eye migration (Keefe and Able
            1994). Many were capable of partial burial at early stages (stages G to H-, Fig. 2.6),
            but were not able to bury completely until late transformation (H+). Burke (1991)
            suggests that these behaviors may begin to develop during transformation offshore.










             Chapter 6
             Estuarine Juveniles


             INTRODUCTION


             Permanent settlement and subsequent fast growth of juveniles occurs in estuarine
             nursery grounds, and recent evaluation of age data has determined that juveniles may
             become sexually mature as early as the end of their first summers (Able et al. 1990;
             Szedimayer et al. 1992; Almeida et al. 1992). The temporal occurrence of juveniles in
             estuaries varies with location. In the northern part of their range, they leave during the
             winter and return the next spring with the adults. In the southern part of their range,
             they enter estuaries in the winter and may stay until the following winter.

             SPATIAL AND TEMPORAL DISTRIBUTION


             The best available information indicates that juveniles occur most frequently in shallow
             subtidal and intertidal areas in the lower portions of estuaries (Tagatz and Dudley
             1961; Keup and Bayless 1964; Dahlberg 1972; Powell and Schwartz 1977), but the
             habitats may change with increasing size. In the South Atlantic Bight, recently settled
             individuals (< 25 mm TQ occur on tidal flats and in marsh creeks (Powell and
             Schwartz 1977; Weinstein and Brooks 1983; Burke et al. 1991). In Charleston Harbor
             (Fig. 2.7b), recently settled juveniles (< 25 mm TQ occurred in a number of estuarine
             creeks (Fig. 6.1) in the winter (January/March), which is consistent with the winter
             occurrence of larvae in this region (McGovern and Wenner 1990; Warlen and Burke
             1990). Less is known for the Middle Atlantic Bight. In New Jersey, juveniles less than
             50 mm TL are not frequently collected even though larvae are common (Able et al.
             1990; SzedImayer et al. 1992; Keefe and Able 1992). The juveniles that have been
             collected were in many habitats including salt marsh creeks, shallow coves and
             shallow portions of bays (K.W. Able, unpubl. data). In Virginia, recently settled
             juveniles have been collected in shallow marsh habitats on the western shore of
             Chesapeake Bay and the Atlantic Ocean side of Virginia's eastern shore (Wyanski
             1990).

             Larger juveniles (> 50 mm TL) have been collected in estuaries all along the east
             coast. This has been documented for New Jersey (Szedlmayer et al. 1992; Rountree
             and Able 1992a,b), Delaware (Smith and Daiber 1977; Malloy 1990), Maryland
             (Schwartz 1961a,b), Chesapeake Bay and Virginia's eastern shore (Horwitz 1978; Geer
             et al. 1990; Bonzek et al. 1991; Wyanski 1991; Bonzek et al. 1992), North Carolina








            30
                                                                Estuarine Juveniles





                    Creek          NOVEMBER                        APRIL
                20     Pita           n=O     80                   n  143
                       Lachicotte
                                              40
                lo     Beresf ord
                       Inlet
                 01                           .0
                20                 DECEMBER   20                    MAY
                                      n=O                           n 31


                10                            10

             C0
             _J
                 01

                                    JANUARY                       JUNE
                W                     n = 25  20                   n =87
             z
             LL 10@                           10
             0

                                               0
             UJ  0 ... ..A
                                   FEBRUARY
             M  160                           20                    JULY
                                     n=246                          n=1

                80                            10



                 0
                80                   MARCH    20                  AUGUST
                                     n = 134                        n = 0,


                40                            10
                 012                          oil  30 50 70 90 110 130 150
                   10 30 50  70 go 110 30 150

                                      TOTALLENGTH(mm)
                   Figure 6.1. Length -frequency distributions for juvenile summer flounder collected
                   during 1986 (November, December) -1987 from four Charleston Harbor estuarine
                   system marsh creeks (Fig. 2.713) using a rotenone/block net method (Wenner et
                   al. 1990).
                   Ll






              Estuarine Juveniles
                                                                                                     31


              (Burke et al. 1991), South Carolina (Wenner et al. 1990) and Georgia (Reichert and van
              der Veer 1991). Additional details are available from Long Island Sound (Fig. 2.7c)
              where juveniles and adults, approximately 200 - 400 mm TL (Fig. 6.2), were collected
              throughout much of the sound (Fig. 6.3).

              The temporal pattern of estuarine use by these larger juveniles varies with latitude and
              presumably winter water temperatures. In Long Island Sound, they appear most
              abundant in spring and fall, and are less abundant in mid-summer (Fig. 6.2, 6.3), when
              they may have moved into subestuaries of the sound. By late fall/early winter
              (November), they have moved offshore into the Atlantic Ocean. In Great Bay, young-
              of-the-year spend the summer in the estuary and begin leaving as early as August
              (Able et al. 1990; Rountree and Able 1992a; Szedimayer and Able 1992; SzedImayer et
              al. 1992) and this continues until November or December, when very few individuals
              remain. In Delaware Bay, most individuals were collected from May through
              September but a few were taken in every winter month in the deeper parts of the
              estuary (Smith and Daiber 1977). In Virginia waters of Chesapeake Bay, most
              collections are dominated by young-of-the-year, at least in most years (Horwitz 1978;
              Geer et al. 1990; Bonzek et al. 1991; Wyanski 1990; Bonzek et al. 1992). These
              individuals tend to be closer to the ocean in March and are more abundant farther up
              the bay during the summer and fall. Typically, all juveniles and adults move offshore
              for the winter (Hildebrand and Schroeder 1928; Musick 1972), although this pattern
              may vary between years. In a warm year (1991) in Chesapeake Bay, some individuals
              were captured during January while in other years (1989, 1990) they were absent
              (Horwitz 1978; Geer et al. 1990; Wyanski 1990; Bonzek et al. 1991, 1992).

              For Pamlico Sound and the adjacent subestuaries in North Carolina (Fig. 2.7d),
              juveniles are most abundant in the summer but are collected as late as December
              (Fig. 6.4) and reportedly remain in the estuaries for the first 18 to 20 months of life
              (Powell and Schwartz 1977). The earliest collections available (March; Fig. 6.4)
              potentially represent individuals that overwintered. By May, the year class that
              entered as larvae in the winter (see above) is evident at sizes of 20 - 100 mm TL   in
              shallow beach and marsh fringe habitats (Fig. 6.5). This same cohort is evident in
              trawl collections from the deeper portions of Pamlico Sound in seasonal collections
              from June through December (Fig. 6.4). These individuals are most abundant in the
              central portion of Pamlico Sound and are seldom found in the larger subestuaries
              (Pamlico and Neuse rivers) in the system, regardless of whether it is a year of
              relatively large (1987) or small (1990) abundance (Fig. 6.5). A seasonal pattern is
              evident with low abundance in the spring (March) and winter, and greatest abundance
              in summer (June and September) (Fig. 6.6). In Charleston Harbor (Fig. 2.7b), larger
              juveniles are evident in the harbor channels during bimonthly sampling (Fig. 6.7), thus
              at this latitude, a portion of the population overwinters in the estuary. In Georgia,
              juveniles have been collected from March through July and are most abundant in April
              and May (Mahood et al. 1974a,b,c,d); Shipman 1983; Music and Pafford 1984).








                        32
                                                                                                                                       Estuarine Juveniles


                                                      20                 1987                                          1990
                                                                        n   526                                       n = 243
                                                      16                                 APRIIL                                        APRIL
                                                      12                                 n = 93@                                       n  19
                                                      8

                                                      4

                                                      0
                                                      20
                                                      16                                 MAY                                           MAY
                                                      12                                 n = 82                                        n = 41

                                                      8

                                                      4
                                                      0                                                                                PL_ A
                                                      20
                                                      16                                 JUNEL                                         JUNE
                                                      12                                 n=27                                          n=7

                                                      8

                                                      4
                                                      0                 1 A-1.      .... . .
                                           U)         20
                                           _J
                                                      16                                 JULY                                          JULY
                                                      12                                 n = 39                                        n=5


                                                      4
                                           z          0        P 11141        UP ... Imp ... 11.1 ... C.           oil
                                           LL         20
                                           0          16                               AUGUST                                          AUGUST
                                           cc
                                           LLI        12                                 n  86                                         n = 66
                                           00         8
                                           2
                                                      4
                                           z          0                 111111-Al  &1    4,     1 IL                                   a
                                                      20
                                                      16                            SEPTEMBER                                          SEPTEMBER
                                                      12                                 n = 70                                        n = 49

                                                      8

                                                      4
                                                      0             111 111111111 AIJI 18110                            1116111   ---------
                                                      20
                                                      16                             OCTOBER                                           OCTOBER
                                                      12                                 n  101                                        n = 56

                                                      8

                                                      4
                                                      0             1, Id                      0-                 A       -1-11
                                                      20
                                                      16                             NOV EMBER                                         NOVEMBER
                                                      12                                 n=28                                          n=1

                                                      8
                                                      4                 ...
                                                      0---..
                                                      R    z                                             ZZ,         'Z' -p    p,
                                                          IC, lp 5@'    4" 49 4 "N
                                                                                                                              0


                                                                                    TOTALLENGTH(mm)
                                                      Figure 6.2. Monthly length-frequency distributions for juvenile and                     adult
                                                      summer flounder in Long Island Sound (Fig. 2.7C) during years of high (1987)
                                                      and low (1990) abundance. Collections were made with 0.5 hour tows of a 14
                                                      m otter trawl at, typically, 40 stations that were chosen by stratified random
                                                      design. Data are based on the finfish surveys of the Connecticut Division of
                                                      Marine Fisheries (1 990a, 1990 and 1992).
                                                      A
                                                      L4






                            Estuarine Juveniles
                                                                                                                                                                                                                          33





                                                                                             1987                                                                     1990

                                                                                                                                   APRIL
                                                                                                                                                                                     ................
                                                                                                                                                                                                              0
                                                                                             -so
                                                                                                                                   n        18                                             6       00 0
                                                                                                           0
                                                                                                                                                                      0 0            so      80
                                                                                                   0.
                                                                                                                                                                           00 0
                                                                                                                                                        0 0       0*

                                                                      0
                                                                                             .. ..........
                                                                                             00


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

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

                                           MAY                                                                                       MAY
                                                                                                                     0             n        43                                       .:.,o           0
                                                                                                                                               ....... . .....
                                                                                                                                               . . ...... ...
                                                                                                                                               .... ... . .
                                                                                                                                                                                     0
                                                                                                               0
                                                                                                                                               ........ . . ....
                                                                                                                                               . ............. ....
                                                                                                                                               .. . ..... .
                                                                                                                                                                                           0
                                                                                             00:0                                                                          00
                                                                                                                                                        0
                                                                                                                                                                                           8*
                                                                                                                                                                      go             0
                                        X,
                                                                                                                                            0
                                                                                             ...............
                                                                  . ......                   ..... ...................... ..........
                                                                  ............. .............. .....
                                                                      .......                ....i............. ...
                                                                  . ........                 .. . ..... ..
                                                                                                                                                            ............. ........
                                                      . .... ..
                                                     ... . ....       ........ ..... 1 11 11 .......... ..
                                                                  ........                   . ....................
                                                           .....  . .......... .             -..........................
                                                                  .. ... ............ ................
                                                                  ...... ...............     ..... ..... .
                                                                                             ...... . ..... ......... ...
                                                                                                                                                                  ........ ......... .
                                                                  . . ............. .......                                                            ...... . .. .............
                                                                                             ... ..... ...
                                                                                                                                                                  ...............
                                                             .... .....................      .. . ..
                                                                      ..........             -- .....                                                                 .. 11 ....
                                                                  . .................        ..
                                                                                                                                                                  ... ........

                                                                                                                                    JUNE                          .....                                    60
                                                                      ............                                                  n=7
                                           n   27                                                          0
                                                                                                                                                                                           0     0
                                        ....                                                 O's                                                                                     0                 0
                                        ....                                                         00                                                                                0
                                                                                                                                                                                     0   0       0
                                                                      0                      08 0000
                                                                                                                                                                                  0
                                                                                                                                                                      0
                                                                      0                      0                                                                                         0
                                                                                             0   0                                                                                   0       0
                                                                                                                                                           0000
                                                                  0                                                                                                            Oo      0
                                                                                                                                                          0       8
                                                                                                                                                                                     0
                                                                      0
                                                                                             0                                                                                                             .......
                                                                                                           .. .....                                                   ..
                                                                                                                                            0
                                                                                                                                            0


                                                                                                                                     .. .......        ... ....   .....
                                                                                                                                                                               .........
                                                                                                                                                                               ...........
                                                                                                                                                                  ...... .. .... .I,:



                                                                                                 a                                          5
                                                                                             .0000                                                                ........
                                                                                                           0                                          .......                                    0
                                           .........                                         0   000
                                                                                             0                                                                             0
                                                                                                 0
                                                                                             so
                                                                                                                                                                      0
                                                                                             0go
                                                                  O@                                                                                      0       0                                        NUMBER
                                                                      00
                                                                      0
                                                                                                                                                                  0                                        OF FISH
                                                                                                                                                          0                       0
                                                                                             .......                                                                                             SYMBOL I TOW
                                           ......         ... ...
                                                                                                                                                                                                     0
                                                                                                                                                                                                     0

                                                        . ... .......
                                                          ...........-
                                                                                                                                                                                 ... ..                        -20


                                        Figure 6.3. Distributio                              n and abundance of juVenile and adult summer flounder in
                                        Long Island Sound (Fig. 2.7C) during years of high (1987) and low (1990)
                                        abundance. * Collections were made with 0.5 hour tows of a 14 m otter trawl at,
                                        typically, 40 stations that were chosen by stratified random design. Data based on
                                        the finfish surveys of the Connecticut Division of Marine Fisheries (1 990a, 1990b and
                                        1992).








                                               34
                                                                                                                                                                                                                                                  Estuarine Juveniles




                                                                                                                  1987                                                                                                               1990

                                                                  ...........
                                                               FAUGUST
                                                                                                                                                                 0                                                                                                           000
                                                                     n=86                                                                             0                                                                            X- ... X

                                                                                                                                         0
                                                                                                                      00                            0
                                                                                                                                                                                                                                                         0
                                                                                                                      0     80           0
                                                                                                           0
                                                                                                  0     0

                                                                                 0
                                                                                                                                   . ............ .   Xx.
                                                                                                                              ......................
                                                                                                                             ........................
                                                                                                                         ... ..................
                                                                                                                      ........................                                                                                      .. ........ .
                                                                                                                            .................. .
                                                                                                                                                                                                                                          .........      ......
                                                                                                                         ........... ..........
                                                                                                ... .                          ....                                                                                               . ......


                                                                                                                                                                                              ........................
                                                                                                                                                                                         SEPTEMBER
                                                                  SEPTEMBER
                                                                         n       70                                                                   0                                             n=49
                                                                                                                                                                                                                                                                             08
                                                                                                                                                                                                                                                                                     0
                                                                                                                                                                                                                                                         0
                                                                                                                                         00                                                                                                                       0    00
                                                                                                                                                    0
                                                                                                                        00                                                                                                                  988
                                                                                                                                                                                                                                          0
                                                                                               0
                                                                                                                                                                                                                         0
                                                                                                                                                                                                                                                                             . . .. .......
                                                                                                                                                                                                                                                                                    ..........


                                                                                                                                                                                                    ............
                                                                                                                                                                                                    ............
                                                                                                                                                                                                    ............
                                                                                                                                                                                                    ...........
                                                                                                                                                                                                    .......... ..
                                                                                                                                                                                                    . ...... 1 11
                                                                                                                                                                                                    . ...... . ..
                                                                                                                                                                                                    ........... ...

                                                                                                                                                                                ... .....................................................
                                                                    OCTOBER
                                                                                                                                                            ........    ... 0
                                                                                                                                                                                           OCTOBER

                                                                                                                                                                                                                                                         0.
                                                                       n       101
                                                                                                                                                                                                    n  53
                                                                                                              ... ..                                                                                                  ...... I
                                                                                                                                                                                                                                                  0               0
                                                                                                                                                    0
                                                                                                                                                                                                                                                            0       0        0

                                                                                                  0
                                                                                                                         oleo                                                                         . ...... .                            1     00 0
                                                                                                                                                                                                           .........
                                                                                                                                                                                                                                                            so
                                                                                            0
                                                                                                          0     see 0                                                                                                 000                   0               0
                                                                                                                                                      . .. .....


                                                                                                                  . . . . . . .    . . . . . . . . . . . . . . . . .
                                                                                                          .. . . . I I . . . I I . .     . . I . . ...

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



                                                                                .............                                                                                          .............. .......
                                                                                                                                                                                                                                                                                    :.x
                                                                                                                                                                                                            ER
                                                                                                                                                                                      @JNOVEMB
                                                                 NOVEMBER
                                                                                                                                                                                                                           .................... . 11 1
                                                                         n=2                                                                                                                        n=l               X.
                                                                                   9                                                                     0                                                                                                                   0
                                                                                                                                                                                                                                                                               0        0
                                                                                                                                                                                                                                                                  8
                                                                                                                                                                                                                                                  0
                                                                                                                    0                    0*         8                                                                    ...........
                                                                                                                                                                                                                                                         8      000          0
                                                                                                                                                    00
                                                                                                                            9o
                                                                                                                                                                                                                                                         0
                                                                                                                                         0                                                                                                                             0
                                                                                                                                                                                                                                                  0                 0
                                                                                                                                                                                                                                            0                                             NUMBER
                                                                                                                                                                                                                                                                                          OF FISH
                                                                                                                                                                                                                              0 0         0
                                                                                                                                                                                                                                                         0
                                                                                                  0                                0.
                                                                                                                                         .0                                                                                                                                  -X: SYMBOL /TOW
                                                                                                                                                                                                                                                                                  0
                                                                                                                                                                                                                              ...............
                                                                                       ...........


                                                                                       @:;:-X.:        a
                                                                                                                                                                                                                                                                                                -20


                                                               Figure 6.3 (continued).












                                                                                                                                                                                                                 M
                                                                                                                                                                                                                 U)
                                                    200-,                          1987                                                          1990
                                                    160-,                      n      19644           MARCH                                    n      710
                                                                                                                                                                     MARCH                                       :3
                                                                                                     n     139
                                                    120",                                                                                                            n   23                                      CD
                                                                                                                                                                                                                 C_
                                                     807

                                                     40-                                                                                                                                                         CD

                                                                                                                                             ION . . .
                                              W     208'.         .............
                                                                                                                                                                                                                 CD
                                                    160-'.                                                                                                                                                       W
                                                    120-.                                             JUNE
                                                                                                     n = 920                                                         JUNE
                                                     80-                                                                                                             n   152

                                                     40.
                                              z          -
                                                        0'                                                                         ........           ........
                                              LL    200.
                                              0     160-
                                              cc         7                                        SEPTEMBER
                                              W     120-                                              n = 470                                                   SEPTEMBER
                                                                                                                                                                     n = 535
                                                     80

                                                     40
                                              Z         0                                                                                                                      A
                                                    200

                                                    160
                                                    120                                           DECEMBER                                                       DECEMBER
                                                                                                       n=115                                                     NO SAMPLE
                                                     80

                                                     40

                                                                                                                                                         @p @p       CP
                                                                       fp 5P A 4           @5-     A                                's b,
                                                                     @" A,: k\,       k, X'@ A,! X" N@                             AS: N@ N: AS! A%: XN@
                                                                       NtK NIb N(b 61                                Ib Ib      NC@ NII ,Z N
                                                                                                                              'ZI            00

                                                                                                  TOTALLENGTH(mm)
                                     Figure 6.4. Length-frequency distributions for juvenile and adult summer flounder in the Pamlico-Albe.marle
                                     sounds estuarine system (Fig. 2.7D) during years of high (1987) and low (1990) abundance.                                        A stratified random
                                     design was used to choose sites throughout Pamlico Sound, the lower reaches of Albemarle Sound and three
                                     subestuaries (Pamlico, Neuse and Pungo rivers) for sampling by 20 minute tows of a Mongoose trawl during four
                                     months in 1987 and 3 months in 1990. Data are based on the North Carolina Division of Marine Fisheries
                                     (1987-1991) trawl survey.







                  36
                                                                                                          Estuarine Juveniles



                                    16-
                                    12      E 1987 (n = 185)                                             MAY
                                            E2 1990 (n = 80)                                           1987 n    28
                                     8. ,
                                        -                                                              19,90 n   15
                                     4
                                     0
                                    16-
                                    12.                                                                  JUN
                                     8.                                                                1987 n = 75
                                                                                                       1990 n = 24


                                     0                                                                           ap
                                    16-
                              Cn    12.                                                                  JUL
                                J
                              4      8-                                                                1987 n = 27
                              n                                                                        1990 n = 21
                              In     4.
                                     0                P PAW1 COJIM IN P                     1P P .     W         14
                                    16-.
                              z
                                    12'                                                                  AUG
                                                                                                       1987 n = 20
                              0      8-
                              cc        -                                                              1990 n = 13
                              LLI    47
                              co
                                     0
                                                                                                                  12L
                                    16-
                              z
                                    12.                                                                  SEP
                                     8.                                                                1987 n    19
                                                                                                       1990 n    1
                                     4.



                                    12.                                                                  OCT
                                     8.                                                                1987 n    13
                                                                                                       1990 n    6
                                     4-
                                    . . . . . . . . . .       0 12 P P                         0 ON          0

                                    12.                                                                  NOV
                                                                                                       1987 n = 3
                                     8.                                                                1990 NS
                                     4-
                                     0  ... . . . . . . .


                                                          TOTAL LENGTH MIDPOINT (MM)

                                  Figure 6.5. Monthly length-frequency distributions of juvenile summer flounder
                                  in shallow beach and marsh fringe habitats in the Pamlico-Albemarle sounds
                                  estuarine system (Fig. 2.713) during years of high (1987) and low (1990)
                                  abundance. Sampling was conducted from May through November by the
                                  North Carolina Division of Marine Fisheries with 10.5 and 20 m seines. Data
                                  from Noble and Monroe (1991).







                                         Estuarine Juveniles
                                                                                                                                                                                                                                                                                                      37

                                                                                           1987.                                                                                                                   1990
                                                                                                                                                                         ...............

                                                         MAR
                                                                                                                                                                                     MARCH
                                                                                                                                                                                     n     23
                                                         n            139




                                                                                                                                                                                                                                                                                00



                                                                                                                                                                                                                                                                     so
                                                                                                                                                               . . ..........................



                                                                      .............. .
                                                                                                                                                                                                                                                                 0
                                                                                                                                           0        0  0                                                     0
                                                                                                                                                                                                                                                                   000          0
                                                                                                                                        0
                                                                                                                                                                                                                                          (D

                                                                                                                                                                                                                                 0           0    0



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



                                                                             . . ..... .        .. ..
                                                    ...                  . . ... ...
                                                    ...                  . ...... ...             ....
                                                    ...                            .......
                                                    .........            ................
                                                                                                                                                                                              ..... .. ...      .......
                                                                                                                                                                                     .......... .....      .. ........
                                                                                                                                                                                     .................   ...
                                                                                                                                                                                                 . ...... ...... .
                                                                                                                                                                                     ................
                                                                                                                                                                                     .. ...... .               .....
                                                                                                                                                                                     .... .... 1 11 .
                                                                                                                                                                                     . . .........
                                                                                                                                                                                     .. ........ ....
                                                    . . . . . . . . ...... ....... ...                                                                                                     ........ . .
                                                                                                                                                                            . ....... .




                                                                                           .. ........
                                                                                                                                                                                     JUNE
                                                      in         1    195                                                             0                                                                                          . ............
                                                                                                                                                                          il         n    214          r

                                                                                                                                                 ih




                                                                                                                                                                                                                                                                        00
                                                                                                                              j:j     G)
                                                                                                                                                         ON;,
                                                                                                                                                         Q)


                                                                                                          .............
                                                                                                         ......  ......
                                                                                                                                               %
                                                                                                      Q                                                                                                                          %
                                                                                                                                                                           .......... .........
                                                                                                                                                                                                   ........     ...
                                                            ..........I...............
                                                                                                                                                                                                                             Go


                                                                                                                                                                                                                                                                                NUMBER
                                                                                                                                                                                                                                                                                OF FISH
                                                                                                                                                                                                                                                       SYMBOL                   TOW
                                                                                     . .. .. .......                                                                                                          .... ..   -

                                                    .. .............. ................     .....
                                                                                                                                                                                     . . . . . . . .. . . . . . . .. . . . . . . .                            0                 0
                                                                                                                                                                                            ....................  -   ......
                                                                                                                                                                                                            . ........ .. .
                                                                                                                                                                          . . . . . . . . . . . . . . . . . .
                                                                                                                                                                                                                                                              0                 1-5
                                                    ....... ...
                                                      ... . . ..... .
                                                                                                                                                                                                                                                                                -10
                                                                                                                                                                                     ......... .... .....                                                                       6
                                                                      . . ....                                                                                                                                                                                                  -20
                                                        ....          .. ...
                                                                                                                                                                                                                                                             Q                  21-50
                                                                                                                                                                                                                                                                                51-100


                                                Figure 6.6.                         Distribution and abundance of juvenile and adult summer flounder                                                                                                                            in the
                                                Pamlico-Albemarle sounds estuarine system (Fig. 2.7D) during years of high (1987) and low (1990)
                                                abundance. A stratified random design was used to choose sites throughout Pamlico Sound, the
                                                lower reaches of Albermarle Sound and three subestuaries (Pamlico, Neuse and Pungo rivers) for
                                                sampling by 20 minute tows a Mongoose trawl during four months in 1987 and 3 months in 1990.
                                                Data are based on the North Carolina Division of Marine Fisheries (1987-1991) trawl survey.









                                              38
                                                                                                                                                                                                                                                       Estuarine Juveniles




                                                                                                        1987                                                                                                                          1990

                                                                                                                                                                                       ...................
                                                                                                                                                                                                                                    .... ......                            ... ...
                                                              SE TEMBER
                                                                                                                                                                                                                       E
                                                                                                                                                                                           SEPTEMB R
                                                                      n         504
                                                                                                                                                                                                             667






                                                                                                                                                                                                                                                                   ..........



                                                                                                                                                                                                                                                                                   00
                                                                                                                                                                                                                                                                              0

                                                                              .............                                                                          G
                                                                                                                                                                                                                                                                                   40
                                                                         .. ...... .......
                                                                                                                                            S


                                                                                                                                  0                                ...


                                                                                                                                                                                                          ... ......... .
                                                                                                          P2        0                                                                         .... . ......




                                                                                                .....    .........  .
                                                                                                       @X: X        ..:7



                                                                                                                                                                                                       . . . . . . . . . . .


                                                                                                                                                                                    IL

                                                          .................. ............. ... .
                                                         X      DECEMBER
                                                                      n=115

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



                                                                                                                                                              0
                                                                                                                                                                 00
                                                                                                                                                                    %





                                                                                                                                                              0
                                                                                                       0


                                                                                                XX.
                                                                                                                    00
                                                                                                      0     0
                                                                                                    0@:ii;          0,                                        NUMBER
                                                                                                                                                              OF FISH
                                                                                                                                           SYMBOL                 TOW
                                                                                                                                                  0                 0


                                                              .......... ....                   .... ......                                                       640
                                                                                                                                                                11-20
                                                                                                                                                                21-50
                                                                                                                                                               51-100



                                                        Figure 6.6 (continued).
                                                                                                                    ....... .. .
                                                                                                             . ......... ...
                                                                                                         ..... ........

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




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



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







                Estuarine Juvenil          es
                                                                                                                              39



                                                   30                             JANUARY
                                                   20                                n =67

                                                   10



                                                   30                               MARCH
                                                   20                                n = 136

                                                   10-
                                                    0.1
                                               W   30                                MAY
                                                I
                                               <   20                                n = 161

                                                   10


                                                    0
                                               AL
                                               0
                                               M   30
                                               W                                     JULY
                                               M                                     n = 162
                                               2   20
                                               M
                                               z
                                                   10


                                                    0
                                                   30                           SEPTEMBER
                                                   20                                n = 23

                                                   10


                                                    0
                                                   30                            NOVEMBER
                                                   20                                n = 74

                                                   10


                                                    01
                                                          4         12        20         28
                                                                 TOTAL LENGTH (cm)
                                    Figure 6.7. Length-frequency distributions for juvenile summer flounder
                                    in channels in the Charleston Harbor. estuarine system (Fig. 2.7B)
                                    during 1985-1988. Bimonthly sampling was conducted using 4.9 and
                                    7.6 serniballoon shrimp trawls with 3.2 - 32 mm mesh liners in the
                                    codends. Data based on a survey conducted by the South Carolina
                                    Marine Resources Center (C. Wenner, pers. comm.)








               40
                                                                                    Estuarine Juveniles


               TEMPERATURE, SALINITY AND DISSOLVED OXYGEN

               Estuarine juveniles can be found in a wide range of naturally occurringr physical
               conditions. Most laboratory and field studies are in agreement regarding the
               preference for higher salinity, at least at larger sizes. In the winter, collections of
               recently settled individuals (< 50 cm TL) in the Charleston Harbor estuary (Fig. 2.7b;
               Table 6.1) occurred at very low as well as high salinities (February - March; Fig. 6.8).
               However, by May, when most individuals ranged from 20 - 110 mm TL, they were
               found at higher salinities (> 10 ppt). Thus in this system, as they disperse into the
               estuary, they may move up into nearly fresh water, but as they grow they are most
               abundant in the higher salinities that occur lower in the estuaries. In lower
               Chesapeake Bay, young-of-the-year were common in creeks that had salinities over
               15 ppt and were most abundant at the highest salinities but were absent in a creek
               where values were 3 - 11 ppt (Table 6.2). In North Carolina, young-of-the-year have
               been found at salinities from 3 - 35 ppt, but were most abundant when salinities were
               greater than 12 ppt (Powell and Schwartz 1977). In more recent data from Pamlico
               Sound (Fig. 2.7d), almost all individuals were collected in the sound while few were
               found in the adjacent lower-salinity subestuaries such as the Pamlico and Neuse
               rivers (Fig. 6.6). This pattern was similar in years with both high and low abundance.

               A number of laboratory studies indicate that juveniles grow best in moderate
               temperatures and higher salinities. For example, feeding rate is positively correlated
               with temperature but interacts with salinity so that higher salinities result in faster
               feeding rates (Peters and Kjelson 1975). These studies found that weight gain and
               salinity were positively correlated between 10 - 30 ppt but reduced at 40 ppt. Other
               studies (Malloy and Targett 1991) found little effect of salinities (10, 20 and 30 ppt) on
               feeding rate, assimilation efficiency and growth rate. Studies designed to test the
               effects of temperature on condition, growth and survival of juveniles from estuaries
               north (Delaware) and south of Cape Hatteras (North Carolina) indicate that differences
               in late winter/early spring water temperatures have significant effects on growth
               (Malloy 1990; Malloy and Targett 1991; Malloy and Targett 1994; Malloy and Targett in
               press). Southern juveniles generally experience warmer temperatures in the field and
               showed increased vulnerability (reduced growth and increased mortality) to cold water
               conditions in the laboratory compared with northern juveniles.

               Low growth rates have been observed at low temperatures (< 12 - 140C) in the,
               laboratory (Malloy 1990; Malloy and Targett 1991; Malloy and Targett 1994; Malloy and
               Targett in press) and have been confirmed in caging experiments in New Jersey (Fig.
               6.9). During the autumn, the average growth rate was negligible (-0.6 and 0.01
               mm/day). Under these same conditions there was little change in developmental











                                                                                                                                                                                       M
                                                                                                                                                                                       (n
                                                                                                                                                                                       C


                                   Table 6.1. Physical characteristics of four marsh creek study sites in the Charleston Harbor estuary (Fig. 6.1) sampled monthly                     CD
                                   for 13 months from June 1986 - August 1987 (Hoffman 1991). Fishes were collected by setting paired block nets upstream                              C_
                                                                                                                                                                                       c
                                   and downstream of the sites, releasing rotenone upstream, and then dip-netting and seining. See Fig. 2.713 for station                              <
                                   locations; 1 = Pita Creek; 2   Lachicotte Creek; 3 = Beresford Creek; 4 = Inlet Creek. (mlw = mean low water.)                                      CD
                                                                                                                                                                                       =3

                                                                                                                                                                                       CD
                                   (SITE NO.)                TEMPER-         SALINITY         DISSOLVED            DEPTH                                    TOTAL NO.                  U)
                                   LOCATION                   ATURE             (Ppt)          OXYGEN                (M)             SUBSTRATE               SUMMER
                                                               (0 C)                             (PPM)                                                      FLOUNDER

                                   (1) 35.6 km from            7.6-32         0.8-24.5          4.2-10.2       0.6 - 1.1 at mIw         shell hash                50
                                   mouth of Charleston                                                                                  and sand
                                   Harbor





                                   (2) 30.7 km from           7.3-30.9        7.1-25.2          1.6-11.2       0.7 - 1.8 at mIw         sand and shell            133
                                   mouth of Charleston                                                                                  hash with some
                                   Harbor                                                                                               mud downstream

                                   (3) 20.7 km from           9.6-31.5       13.0-23.7          2.0-11.7       0.4 at mlw               mud upstream,            223
                                   mouth of Charleston                                                                                  sand and some
                                   Harbor                                                                                               shell downstream

                                   (4) 8.3 km NE of           7.2-32.0       27.4-36.2          0.8-10.7       0.7 at mlw               shell hash with           201
                                   mouth of Charleston                                                                                  very little mud, 2
                                   Harbor in a creek off                                                                                small oyster bars
                                   the Intracoastal                                                                                     in upper 1/2 part-
                                   waterway                                                                                             ially exposed at
                                                                                                                                        low tide









                     42
                                                                                                                        Estuarine Juveniles





                                                                80     Creek                       JANUARY
                                                                60      M Pita                        n=18
                                                                40          Lachicotte
                                                                            Beresford
                                                                20      ED Inlet

                                                                  W
                                                                80                 .7,
                                                                                                   FEBRUARY
                                                                60                                    n = 246

                                                                40

                                                                20

                                                                  0
                                                                80                                    MARCH
                                                                60                                    n = 134

                                                                40

                                                                20

                                                                  0
                                                                80                                    APRIL
                                                           z    60                                    n = 143
                                                           U-   40
                                                           0
                                                           CC   20
                                                           LU

                                                                80"                                   MAY
                                                           z        7
                                                                W                                     n = 31

                                                                40'

                                                                20'

                                                                  0
                                                                80
                                                                                                      JUNE
                                                                60                                    n=6

                                                                40

                                                                20

                                                                  0'
                                                                80                                    JULY
                                                                60                                    n=1

                                                                40

                                                                20
                                                                  01                                  1!9-03
                                                                    N9                                @'- "\.
                                                                  Qr tr V                             fp
                                                                                          lt@      9@1
                                                                                 SALINITY (ppt)
                                                      Figure 6.8. Abundance of juvenile summer flounder
                                                      ,relative to salinity in four Charleston Harbor estuarine
                                                      system marsh creeks (Fig. 2.713) collected by a
                                                      rotenone / block net method during 1987 by the South
                                                      Carolina Marine Resources Center. Data based on
                                                      Wenner et al. (1990).










                                                                                                                                                                                                     M


                                     Table 6.2. Physical characteristics of four marsh creek study sites in lower Chesapeake Bay and the ocean side of the eastern
                                     shore of Virginia (Fig. 2.1) sampled monthly from June - November in 1989 and May - November in 1990 (Daniel in prep).
                                     Fishes were collected by setting paired block nets upstream and downstream of the sites, releasing rotenone upstream, and                                       CD
                                                                                                                                                                                                     C_
                                     then dip-netting and seining.                                                                                                                                   C:
                                                                                                            MEAN                                                                                     CD
                                     (SITE NO.)                        TEMPER-           SALNITY            LOW                                                         TOTAL NO.                    =3
                                                                         ATURE                                            SUBSTRATE             VEGETATION               SUMMER                      CD
                                     LOCATION                                               (Ppt)          WATER                                                                                     CA
                                                                          (0 C)                              (m)                                                        FLOUNDER

                                     (1) small tributary                  15-31             3-11            0.3-0.5       shell hash and          off Spartina                0
                                     of the Poropotank River;                                                             very muddy              marsh, some
                                     -3.1-6.1 m wide, -30.5 m                                                                                     Juncus
                                     long


                                     (2) in Goodwin Islands               16-30             15-24            0.9          hard-packed mud         off Sp-artina               26
                                     at mouth of York River;                                                              and shell hash in       marsh; Spartina
                                     -9.1 rn wide, -30.5 rn long                                                          center, very            island in
                                                                                                                          muddy sides             center



                                     (3) creek off main channel
                                     entering town of Wach-               15-30             26-29           0.8-0.9       shell hash in cen-      none                        59
                                     apreague; -6.1 m wide,                                                 (hole=        ter and steep
                                     -30.5 m long                                                          1.1-1.2)       muddy sides

                                     (4) behind Parramore
                                     Island, close to mouth
                                     of Little Machipongo                 17-28             28-33          0.15-0.2       sandy mud               off Spartina                68
                                     Inlet; -6.1 m wide, -30.5                                           (hole=1.2)                               marsh
                                     m long









                        44
                                                                                                                                Estuarine Juveniles




                                                   12-                                                 n     17
                                                                                                             -::-                    n =7
                                                                     SPRING
                                                                    (MAY-JUNE)
                                                   10

                                                                        n      13
                                                                                                             .........                     ......
                                                     8-                                                      . ... . ...
                                                                                                             ..........                  ... ..
                                                          n      23
                                                                F771                                                                     -:X-X:X-
                                                     6-                        .........
                                             Cl)                                      n =22
                                             E
                                                                 @:X::.X                                                                 ........
                                                                  .......      ....
                                                                    ...        ........       ...
                                             E       4-                                                              n'=
                                                                  .......      ...  ....      .....
                                                                 ........      .........
                                                                                                                         rr77M
                                                                                                                          .........      ..... ...
                                                     2-
                                                                 . .....       .......          ....         ......
                                             0
                                             z                                                                                           .......
                                             LU      0          ....... .
                                             _J           TRIAL I        TRIA 11        TRI     I      TRI@L 11       TRIAL I        TRIAL 11

                                                   12-
                                             UJ
                                             (0                                                                           HABITAT
                                             4     10-
                                             LU                                                                  VEGETATED
                                             W
                                             0       8-             AUTUMN                                                 EELGRASS
                                             z                                                                            SEA LETTUCE
                                                                   (NOVEMBER-
                                             z                      DECEMBER)                                    UNVEGETATED
                                             -4      6-                                                                    MUD
                                             UJ

                                                     4-
                                                                                                       n     4
                                                     2-                                  n    3                                      n     6
                                                                                       loom                  X:
                                                                                                                                         X ......
                                                            n=20 n=18
                                                                                                             ........ no data
                                                     0-
                                                           TRIAL I        TRIAL 11      TRIAL I        TRIAL 11      TRI'AL I        TRIAL 11
                                                         MARSH ELDER                      LITTLE BAY                    SCHOONER
                                                                ISLAND                                                     CREEK
                                                                                           STATION


                                             Figure 6.9. Habitat-specific growth of juvenile summer flounder held in
                                             cages (6 mm mesh) at three sites in the Great Bay-Little Egg Harbor
                                             estuarine system (Fig. 2.7A) during spring and autumn 1991. Two trials of
                                             10 - 11 days duration were conducted at each station during each season.
                                             For each trial, four cages containing juvenile summer flounder (n = number
                                             of individuals used in each trial) of 12 - 41 mm SL were deployed: two on
                                             vegetated substrate (eelgrass = Zostera marina; sea lettuce = Ulva
                                             lactuca) and two on adjacent unvegetated substrate (mud). Adapted from
                                             Keefe and Able (1992).






            Estuarine Juveniles
                                                                                               45


            stage as had been observed in laboratory experiments (Keefe and Able 1994). At
            temperatures below 2 - 30C, significant mortality has been reported in laboratory
            experiments (Malloy 1990; Malloy and Targett 1991; SzedImayer et al. 1992; Malloy and
            Targett 1994; Malloy and Targett in press) and this may be an important source of
            mortality affecting subsequent year-class strength. The impact of low temperatures
            may vary between Years depending on the severity of winter. During spring and
            summer, growth rates have varied from approximately 0.5 - > 1.0 mm/day for recently
            settled individuals (Keefe and'Able 1992) to a range of 1.5 - 1.9 mm/day for young-of-
            the-year juveniles (SzedImAyer et al. 1992; Rountree and Able 1992a). Estuarine
            juveniles with ultrasonic tags that were tracked for 1 - 33 days in Schooner Creek in
            Great Bay (Fig. 2.7a) were observed over a wide range of temperatures (16.0 -
            28.OOC), salinities (22 - 35 ppt) and oxygen levels (2.4 - 8.9 ppm), but they generally
            stayed within narrow limits for these parameters (Szedlmayer and Able 1993). The
            mean values for each fish were much more restricted for temperature (22.3 - 24.90C),
            salinity (27 - 31 ppt) and dissolved oxygen (5.9 - 6.8 ppm).

            SUBSTRATE

            Recently settled individuals have been collected in estuaries on the eastern shore of
            Virginia in depths typically less than 2 m where the substrate was composed of more
            than 50% very fine sand, silt and clay (Wyanski 1990). In North Carolina estuaries,
            similar-sized individuals were abundant in shallow (< 1 m) areas but were also found
            in slightly deeper areas (1.5 - 3 m). They were most abundant over sandy substrates
            (Burke et al. 1991). In laboratory experiments with the same population, juveniles (14.7
            mm ï¿½ 1.2 mm SL) preferred sand substrate (< 5111o silt-clay) over mud substrate (> 95%
            silt-clay) regardless of availability of prey (Burke 1991). This same preference for
            sand was observed for similar stages in the laboratory for a New Jersey population
            (Keefe and Able 1992). In marsh creeks in Charleston Harbor (Fig. 2.7b), similar-sized
            individuals were abundant over substrates that ranged from mud to sand and shell
            hash with occasional oyster bars (Table 6.1).

            Larger young-of-the-year in North Carolina have been most abundantly collected
            where sand sediments or a transition from fine sand to silt and clay occurred and less
            abundantly where silt and clay predominated (Turner and Johnson 1973; Powell and
            Schwartz 1977). They also occurred abundantly in marsh creeks with soft mud
            bottoms and some shell hash in southern New Jersey (Szedlmayer et al. 1992;
            Rountree and Able 1992a), Virginia (Table 6.2) and South Carolina (Table 6.1). In
            Virginia, these larger individuals occurred over shallow sand, deep sand and deep fine
            sediments (Wyanski 1990) as well as in eelgrass beds (Orth and Heck 1980; Lascara
            1981; Weinstein and Brooks 1983).

            Habitat quality, as measured by relative growth, was evaluated with caging
            experiments in Great Bay-Little Egg Harbor (Fig. 2.7a; Fig. 6.9). Growth of recently








               46
                                                                                    Estuarine Juveniles


               settled and small juveniles (17 - 41 mm SQ based on caging experiments was variable in
               spring (range 0.18 - 0.89 mm/day). Growth did not appear to be strictly related to the
               habitats tested (eelgrass and adjacent unvegetated substrate, sea lettuce and adjacent
               unvegetated substrate and marsh creek). The fastest growth occurred in shallow bays
               and marsh creeks.


               BEHAVIOR

               Tagging studies in Long Island (Poole 1962), New Jersey (Hamer and Lux          1962; Murawski
               1970a) and Maryland (Jesien et al. 1992) estuaries indicate that a large proportion of
               juveniles tagged in estuaries in the summer return to the same system during the
               following summer. Diel movements of estuarine juveniles have been studied- in two
               habitat types, marsh creeks and eelgrass habitats, and in the laboratory with sand and
               mud substrates. In the laboratory, recently settled juveniles exhibited a diel pattern of
               burying behavior that was influenced by several other variables as well, including
               substrate type, water temperature,   tide and the presence and type of predator (fish or
               shrimp; Keefe and Able 1994). These individuals buried more during the day, especially
               at the time of high tide. In Great Bay,- ultrasonically tagged individuals followed a regular
               diel cycle of movements in the 1-km-long Schooner Creek (Fig 2.7a; SzedImayer and
               Able 1993). These young-of-the-year individuals (210 - 254 mm TQ.spent most of the
               time at the mouth of the creek during the July-September study period. Movements up
               the creek typically occurred on night flood tides followed by a return down the creek on
               the following ebb tide. These tide-mediated movements.were for feeding on resident
               marsh creek fishes and crustaceans (Rountree and Able 1992a) and may have been
               influenced by low dissolved oxygen conditions in the upper portion of the creek, especially
               on night low, tides (Szedlmayer and Able 1993). In North Carolina, some individuals move
               onto the surface of regularly flooded salt marshes during flood tides (Hettler 1989). In
               both field observations in Chesapeake Bay and laboratory experiments (Lascara .1981),
               juveniles fed in and at the edge of patchy seagrasses (Zostera marina and Ruppia
               maritima) while displaying an ambush predator strategy. In these habitats most feeding
               occurred in the-morning.

               Seasonal movements of larger juveniles      from estuaries occur, as temperatures are
               dropping -in aut'umn but this pattern may vary with latitude, water depth of the estuary and
               temperature. - In-Great Bay (Fig. 2.7a), juveniles migrated out of marsh systems in the late
               summer and early fall. In one, instance, a single individual was tracked from Schooner
               creek, through Little Egg Inlet and into the ocean, a distance of about 1.5 km (Szedlmayer
               and Able 1993). In North Carolina, it has been assumed that juveniles overwinter in the
               estuaries (Powell and Schwartz 1977). However movement offshore, especially of larger
               individuals, may occur in late summer (A. Powell, pers. comm.), which may account for
               the smaller sizes reported for young-of-the-year at the end of the year (Powell 1982)
               relative to more northern populations. A similar migration offshore with increasing size
               has also been suggested for Georgia populations (Music and Pafford 1984).










             Chapter 7
             Offshore Juveniles


             INTRODUCTION


             Juveniles join adults in an offshore migration, but details are lacking. Some juveniles
             that spend the first summer in estuaries move onto the continental shelf as
             temperatures decline in the fall or as they reach larger sizes. After overwintering on
             the shelf, many move back into estuaries for the second summer. Details of their
             habitats and other aspects of the biology are not we I I-known, because they are not
             frequently differentiated from adults. For purposes of this treatment, offshore
             juveniles are less than 320 mm in the fall, winter and spring.

             SPATIAL AND TEMPORAL DISTRIBUTION

             Juveniles appear to make seasonal migrations similar to those of the adults. They
             leave the estuaries during the summer- or early autumn throughout much of their
             range. In the Middle Atlantic Bight, young-of-the-year have been found exclusively
             on the inner continental shelf in fall surveys (Able et al. 1990) and this pattern was
             evident in more recent surveys as well (Fig. 7.1). In the northern Middle Atlantic Bight,
             most collections were very close to shore while farther south they were found farther
             out on the shelf. In the winter, this year class was distributed off the middle and outer
             continental shelf. from Long Island to Cape Hatteras. By spring.. portions of this same
             year class were still found near the edge of the continental shelf but some individuals
             had already moved back inshore.

             In the South Atlantic Bight (Fig. 7.2), the pattern of movement from the estuaries onto
             the continental shelf may differ in timing. South Atlantic Bight trawl surveys in depths
             of less than 10 m (Beatty et al. 1989; Wenner and Sedberry 1989) indicate that small
             juveniles (100 - 200 mm TL) could be found as early as June and that their numbers
             increased through July and August, when the-modal size was approximately 200 mm
             TL. They were especially abundant at sample locations adjacent to major inlets
             (Beatty et al. 1989; Fig. 7.3). By September through November they were collected








                 48
                                                                                                              Offshore Juveniles







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








                                                           I*K1





                                                           a                     AUTUMN 1991
                                                                                    age = 0
                                                                                    n   178





















                                                                                  WINTER 1992
                                                                                     age
                                                                                     n  627






                                                                           W.




                                                         ..........











                                                                                 SPRING 1992
                                                                                    age= 1
                                                                                    n = 98



                                Figure 7.1. Distribution of summer flounder juveniles (< 320 mm TL; 0 - 1 year of age) from
                                National Marine Fisheries Service (NMFS), Northeast Fisheries Science Center (NEFSC)
                                groundfish surveys for A. autumn, B. winter, and C. spring, 1991 - 1992. Sampling was
                                conducted by stratified random design in the Gulf of Maine to Nova Scotia, on Georges
                                Bank, and in the Middle Atlantic Bight (Fig. 2.1 using 0.5 hour tows of a #36 Yankee trawl
                                with a 12.7 mm mesh liner in the coden . Re 1000 m contour is shown. Collections
                                where no juveniles were caught are omitted. (Adapted from maps provided by G.
                                                            %












































                                Shepherd, National Marine Fisheries Service, Woods Hole Laboratory, Massachusetts).






           Offshore,Juveniles
                                                                                                                    40










                               ..........C   Hatteras
                                         ape
                                      Pamlico Sound


                    ..... . ......
                                      ape Lookout
                                                                                    Bogue Inlet
                                    River


                                 ape Fear
                                   Fear River
                                                                                Lockwood Folly Inlet

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

                   ..................
                                 Bay
                                                                             North of Charleston Harbor
                                                                             (index station).
                       ....                                         .. ....
                                                                    .. ......
                           arl ton Harbor                          .........
                                                                                  Inlet   SAMPLING AREA
                           elena Sound
                                                                                           E   Wenner &
                       ort Royal Sound
                                                                                               Sedberry
                    ..,.Savannah River                                                       @7.
                      ORGIA                                                                    Beatty,
                 ...   Itamaha Sound                                                           Webster &
                                                0      100
                                                                                               Wenner 1989

                           Andrews Sound
                             h s River                                WSt Andrews Sound Inlet



                 ..........
                                    Cape C
                                             naveral

                                              River
                    .........                agoon


                Figure  7.2. A. South    Atlantic Bight (region between Cape Hatteras, North Carolina,
                and Cape Canaveral, Florida) with major capes, bays and rivers indicated, and B.
                sample areas for two South Atlantic Bight inner continental shelf (< 10 m) trawl
                surveys. Wenner and Sedberry (1989) sampled at randomly chosen sites within each
                stratum during 1980, 1981 and 1982. Beatty et al. (1989) sampledfour inlet locations
                and an index station during 1987 and 1988. Three sites were sampled at each location;
                a within inlet site, a beach site and an off-beach site.
                                            a








                      50
                                                                                                                              Offshore Juveniles




                                          50   EA. RAND M STATIONS
                                          40  1:113. FIXED INLET          JANUARY                                      JULY
                                                      STATIONS              n = 177                                    n = 302
                                          30
                                                                          (NO SAMPLING
                                          20                              FOR STUDY B) .

                                          10
                                           0                                                                         pip .....

                                          so.
                                          40-                             FEBRUARY.                                  AUGUST
                                                                          (NO SAMPLING                                 n = 554
                                          30@                             FOR EITHER
                                                                          STUDY A OR B)
                                          20.

                                          10.

                                           0

                                          50.
                                                                                                                  SEPTEMBER
                                     (n   40.                               MARCH
                                                                          (NO SAMPLING                                 n = 282
                                          30-                             FOR EITHER
                                                                          STUDY A OR 8)
                                          20.
                                          10'

                                     LL
                                     0    50.
                                     cc   40-                               APRIL
                                                                            n = 185                                 OCTOBER
                                     M    30.
                                                                          (NO SAMPLING                                 n = 280
                                                                          FOR STUDY 8)
                                          20-
                                     z    10.                                                               n
                                           0                           ........

                                          50
                                          40                                 MAY                                  NOVEMBER
                                                                            n = 253                                    n 305
                                          30

                                          20

                                          10

                                           0

                                          50.
                                          40.                               JUNE                                   DECEMBER
                                                                            n=                                         n = 22
                                          30.                                   106                              (NO SAMPLING FOR
                                          20.                                                                      STUDY A: FEW
                                                                                                                 STUDY B SAMPLES)
                                          10-

                                           0                                             .........
                                             10        20         30        40         10         20        30         40
                                                                            TOTAL LENGTH (cm)

                                         Figure 7.3. Monthly length-frequency distributions for juvenile and adult
                                         summer flounder from two trawl surveys conducted in nearshore waters (<
                                         10 m) on the South Atlantic Bight inner continental shelf (Fig. 7.2): A.
                                         random stations sampled between Cape Fear and Cape Canaveral during
                                         1980 - 1982 (Wenner and Sedberry 1989), and B. fixed stations sampled
                                         outside four major inlets between Cape Fear and Georgia during 1987 -
                                         1988 (Beatty, Webster and Wenner 1989).






         Offshore Juveniles
                                                                                                51


         both near the shore and at inlets. During the winter, sampling was less frequent in
         both these programs, but juveniles were abundant near the shore in January. This
         same year class is well represented at these areas in April and May throUgh July and
         August, which indicates that a portion spend the summer on the shallow inner
         continental shelf.


         TEMPERATURE


         In the Middle Atlantic,Bight -collections during 1970-1979, juveniles were most
         abundant at bottom temperatures greater than 159C (Sissenwine et al. 1979), but
         nothing else is known primarily because juveniles and*adults are not differentiated.










              Chapter 8
              Estuarine and Offshore Adults


              INTRODUCTION

              The adult life history stage of summer flounder begins when individuals reach sexual
              maturity. Based on aging and maturation studies (Poole 1961; Eldridge 1962; Smith
              1969; Powell 1974; Smith and Daiber 1977; Morse 1981; Smith et al. 1981; Gillikin and
              Holland unpublished), the majority become sexually mature at sizes as small as 240 -
              270 mm TL for males and 300 - 350 mm TL for females (Morse 1981). The habitats
              occupied by adults are largely determined by their extensive migratory patterns (Figs.
              2.10, 2.11). Details of their seasonal distribution are limited to large-scale trawl
              surveys. These have been more intensive and extensive in the Middle Atlantic Bight
              than in the South Atlantic Bight and this is reflected in the following account in which
              we combine the treatment of adults in estuarine and continental shelf habitats.


              SPATIAL AND TEMPORAL DISTRIBUTION

              In the Middle Atlantic Bight and northern South Atlantic Bight, adults overwinter on the
              outer continental shelf (70 - 155 m depth; Grosslein and Azarovitz 1982), migrate
              inshore to shallow inner continental shelf and estuarine waters in spring, and begin
              offshore migration back to the outer continental shelf in autumn (Figs. 2.10, 2.11;
              Hildebrand and Schroeder 1928; Bigelow and Schroeder 1953; Grosslein and Bowman
              1973; Smith 1973a,b; Rogers and Van Den Avyle 1983; Able et al. 1990) or as early as
              August in some cases (Schwartz 1961a,b). Gonad maturation may begin as early as
              late-summer in Middle Atlantic Bight estuaries (Schwartz 1961a,b; Smith 1969; Smith
              and Daiber 1977), and spawning occurs during offshore migration (Eldridge 1962;
              Smith 1973a,b; Murawski and Festa 1976; Able et al. 1990). There is some evidence to
              suggest that the largest, older individuals eventually remain over the continental shelf
              all year (Festa 1977). However due to current fishing pressure, the majority of adults
              are only three years of age or younger (Terceiro 1993), and those individuals largely
              utilize estuarine and inner continental shelf waters in summer.


              The seasonal patterns of distribution and abundance of adults in the Middle Atlantic
              Bight are largely reflected in the pattern of catches from the commercial fishery (Fig.
              8.1), although these must be cautiously interpreted because the patterns may reflect
              the fisher's behavior rather than that of summer flounder. In January-March when
              water temperatures are coldest, they are clearly most abundant at the edge of the

















                                                                                                                             X ......................
                                   -0-         14
                                                                                                                                      ..............
                                                                                                                                      .............

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



                                                                                                                                                                                                                             ++.+ +
                                                                                                                                                                              . .............. ::::@: ......                    +++
                                                                                                                                  + +
                                                                                                                                +






                                      + +
                                                          NN

                                                                                                                4l'
                                                                          METRIC TONS
                                                                             + 0-10
                                                                             + 10.0-20
                                    +!
                                                                             + >20


                                                                        JANUARY                                                               FEBRUARY                                                                          M






                                                                                                                                                            +++ +                                                 + +


                                                                             ++. +                                                                              Z 11@
                                                                                                                      +


                                                                                                                                                                                                 +
                                                        Aj
                                                 +


                                                                                                                                                                                ...........



                                      ++





                                                                                APRIL                                                                        MAY

                             Figure 8.1. Average annual landings (metric tons per ten minutes square of latitude and longitude) of summer flounder captured in t
                             States commercial fishery during 1987-1989 from the Gulf of Maine to off Cape Hatteras (Fig. 2. 1) by month. The 200 m conto
                                     h  V/* eN
                                                 \0










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







                                ... . . ... ..
                             I







                             (Adapted from maps provided by G. Shepherd, National Marine Fisheries Service, Woods Hole Laboratory, Massachusetts.)















                                                                                                                                                                                        . ..........

                                                                                                        ++   +



                                                                                                                                                                   +  + +


                                                                                                                                                                                                                      ".4
                                      ...........
                                           ..............

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


                                 . .........


                                                                    S%
                                                                                        METRIC @Q@NS
                                                                                           +0_1
                                                                                           + 10.0
                                                                                           + >20

                                                                                                 JULY                                                                    AUGUST,                                                                 SEPI


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





                                                                                                                                                                                                                                                    -++ +
                                                                                                                                                                                                                                             ++.++       +
                                                                               ++.4j:!j@++++ +                                                                  + +       +
                                                                                                                              ....             .......
                                                                             ++ +                                                                                +    ++++4@. . .

                                             .... .......
                                                                                                                                                                                                                                        4.



                                                                                                                                                                                                                  + +

                                                                                                                                                                                                                      4+



                                                                                                                         + +                                                                             +
                                                                                                                                                                                                                A.
                                                                                                                              +                                                                                   4j


                                                                                    OCTOBER                                                                      NOVEMBER                                                                           DE

                                      Figure 8.1 (continued).









              56
                                                                                            Adults


              continental shelf from the westernmost edge of Georges Bank to just south of
              Chesapeake Bay. By April, some catches are reported in shallower waters from off
              Long Island, the south shore of Massachusetts and on to Georges Bank. By May, this
              pattern is more pronounced with catches from inner continental shelf waters from
              Georges Bank south to Maryland. At this time there are much lower catches at the
              edge of the shelf. Catches decline on the shelf through June and July, but they begin
              to increase in August through September. By August, the first reports of catches north
              of Cape Cod occur and these continue through December. Most of the increased
              catches occur off southern Massachusetts and in the vicinity of Delaware Bay. By
              October, this trend continues while spreading over most of the Middle Atlantic Bight.
              In November, catches increase on the middle portions of the shelf, but none are
              reported from Georges Bank. By December, catches at the edge of the shelf have
              increased and the pattern begins to resemble that for January.

              This pattern is supported by the fishery independent data from the long time series of
              trawl surveys conducted by the National Marine Fisheries Service in the Middle
              Atlantic Bight (Fig. 2.10). During the spring survey (February - March) they were most
              abundant at the edge of the shelf from Georges Bank south to the North Carolina -
              Virginia border. This center of abundance is typically in depths of 70 - 150 rn
              (Azarovitz and Grosslein 1987). At the same time, smaller numbers were collected
              over middle to inner shelf waters. More fish were collected in inner shelf waters from
              Delaware Bay and south to Cape Hatteras. In autumn (September - November), fish
              were most abundant from Nantucket Shoals and shallower waters from Long Island to
              the mouth of Chesapeake Bay. Small collections were also made on the mid-shelf
              portions of Georges Bank. Earlier studies, based on trawl surveys prior to 1979
              (Sissenwine et al. 1979), demonstrated a similar pattern with individuals widely
              distributed over the continental shelf in depths from 0 - 360 m during the spring.
              During summer and autumn, they occurred primarily in depths less than 100 m but in
              winter they were not found in water shallower than 70 m.

              The occurrence of adults in estuaries in the Middle Atlantic Bight corresponds in time
              with their inshore movements on the continental shelf. This is evident from several
              systems. In Long Island Sound, individuals that are presumably age 1+ and older (Fig.
              6.2) are present in April and May collections, become less abundant in June and July,
              and more abundant in August through October (Fig. 6.3). Fewer individuals are
              present in November, presumably because of the migration offshore. These are
              distributed throughout Long Island Sound although they are most abundant in the
              central portion. In Great Bay (Fig. 2.7a), based on an extensive creel survey from
              1967 to 1976, adults (> 320 mm TL) are well represented in the catches, particularly
              from 1967 to 1972. Subsequently, the average size of fish landed was smaller during
              1975 to 1976 (Fig. 8.2). In Maryland, adults are present in the estuary as early as April
              and through the summer (Jesien and Hocutt 1991). On the eastern shore of Virginia
              they occur from April - November (Richards and Castagna 1970). In Pamlico Sound







                    Adults
                                                                                                                                        57







                             100-                                             100-1
                              80.                                  1967 80-                                         1972
                              60-.                                             60. '
                              40-                                              40.
                              20.                                              20'
                                0                                                0 "
                             loo,                                             2007
                              80@                                  1968
                              60-
                                                                              100.
                       Cn     40.
                              20
                                0                                                O@
                             loo-                                             200-
                       >                                           1969                                             1974
                              80-
                              60.
                       LL                                                     100
                       0      40.
                       Ix     20.
                       Uj
                                0 .....                                          0                                 ......
                             loo-                                             400
                                                                   1970                                             1975
                       z      80-
                              60 -@                                           200
                              40-
                              20
                                0                                                0
                             100-                                             100-
                              80-                                  1971        80.                                  1976
                              60.                                              60.
                              40-.                                             40-
                              20                                               20 -@
                                0                                                0.
                                    LO   C)   LO   C)   LO    C)   LO C) CO          LO   Co   U")  0    LO   CI    Lo   CD Co
                                    C\I  Co   Co   It   'IT   L0   Lr)   CO(C        N    Co   Co    I-  't    LO   LO   1@0 Co
                                                                            Al                                               Al
                                                                  TOTAL LENGTH (cm)



                             Figure 8.2. Length-frequency distributions of summer flounder in Great Bay (Fig.
                             23A) from the recreational fishery. creel survey conducted by the New Jersey
                             Department of Environmental Protection and Energy for the month of July over a
                             ten year period (Murawski and Festa 1979).
                                                                                     Ali









              58
                                                                                             Adults


              (Fig. 2.7d), adults are notable by their absence (Fig. 6.7) and a similar situation also
              occurs in South Carolina (Wenner et al. 1990; Fig. 6.10) and Georgia (Dahlberg 1972;
              Music and Pafford 1984).

              Information on offshore adults in the southern South Atlantic Bight is limited in part
              because summer flounder are not a major component of the commercial fisheries and
              also because the various species of Paralichthys are not differentiated. There are only
              a few adults (> 300 mm TL) captured during any month in South Atlantic Bight trawl
              survey programs on the inner continental shelf (< 11 m; Beatty et al. 1989; Wenner and
              Sedberry 1989). Prior surveys have captured few adults as well (Struhsaker 1969).
              Adults have been reported on ocean beaches and artificial reef habitats in spring and
              summer in Georgia and Florida (Miller and Jorgenson 1969; Gilmore et al. 1981) and
              on the outer continental shelf in the winter (Pearson 1932).

              TEMPERATURE, SALINITY AND OXYGEN

              Seasonal fluctuations in water temperature may be one of the primary factors
              controlling the timing of spring and winter migrations and spawning for summer
              flounder adults (Nesbit and Neville 1935; Ginsburg 1952; Edwards 1964). Spawning
              adults are found primarily in bottom water temperatures of 12 - 190C (Smith 1973a,b;
              Festa 1974b) on the continental shelf habitat from Cape Cod to Cape Lookout. The
              cold bottom water present in the Middle Atlantic Bight in early autumn may limit the
              seaward extent of migration (Smith 1973a,b), and thus spawning habitat, during part of
              this time. In southern South Atlantic Bight waters, high temperatures may explain the
              absence of adults in estuarine habitats during spring and summer when young-of-
              the-year are present. Most adults occur in the high-salinity (> 28 ppt) portions of
              estuaries (Richards and Castagna 1970; Powell and Schwartz 1977; Mid-Atlantic
              Fishery Management Council 1990; 199ta,b; Burke 1991; Hoffman 1991; Noble and
              Monroe 1991; SzedImayer et al. 1992).

              Summer flounder oxygen requirements have not been well studied, however some
              reports indicate that episodes of hypoxia (low dissolved oxygen, -< 3 ppm) or anoxia
              (absence of dissolved oxygen, 0 ppm) may be a common feature of the offshore
              habitat and may significantly impact local distribution and survival. One of the best-
              studied episodes of hypoxia to affect summer flounder occurred on the Middle Atlantic
              Bight continental shelf off New Jersey in midsummer to late-autumn 1976 (Swanson
              and Sinderman 1979). Reports of dramatically increased catch rates of adults in the
              ocean beach and estuarine recreational fishery indicate that these fishes were
              avoiding hypoxic areas offshore by moving into Great Bay (Murawski and Festa 1977).
              During this event, summer flounder were caught only in ocean areas free of hypoxic
              water, and divers observed dead adults on the ocean floor (Freeman and Turner
              1977). Adult summer flounder were also reported among the victims of a fish kill
              washed onto Jones Beach, New York, in September 1951 during a period of offshore







            Adults
                                                                                    59


            hypoxia similar to the 1976 episode (Perlmutter 1959). Other fish kills reported for
            1968, 1971 and 1974 suggest that mortality due to hypoxia is not uncommon (Freeman
            and Turner 1977).,

            SUBSTRATE

            Adult summer flounder are typically. described as preferring hard, sandy substrate in
            which they can easily bury (Bigelow and Schroeder 1953; Schwartz 1964;.$mith 1969).
            During their stay in Middle Atlantic and northern South Atlantic estuaries, however,
            adults exploit a broad range of lower and mid-estuary, habitats including salt marsh
            creeks (Dahlberg 1972; Rountree and Able 1992a) and seagrass beds (Bigelow and
            Schroede'r 1953; Orth and Heck 1980), which frequently have muddy or silty substrates,
            as well as sand flats (Dahlberg 1972; Gilmore et al. 1981). Adults with normal pigment
            can take on camouflaging patterns that resemble nearly any bottom substrate (Mast
            1916).










              Chapter 9
              Implications for Resource Managers


              Despite the importance of summer flounder to commercial and recreational fisheries,
              we lack a clear understanding of habitat requirements for all life history stages,
              particularly for the eggs and larvae. For instance, we know nothing about the role of
              transport processes on the behavior that influences the movement of transforming
              larvae into estuaries. We know somewhat more for the juveniles and adults, although
              the degree of understanding varies from region to region and is made more difficult
              because this species is highly migratory. In general, our understanding of habitat
              requirements for most life history stages is better for Middle Atlantic Bight than for
              South Atlantic Bight populations.

              Of those habitat parameters for which information is available, it appears that
              temperature and dissolved oxygen have the strongest influence on habitat use and
              quality. Temperature affects the seasonal occurrence in estuarine habitats especially
              in autumn when declining water temperatures result in offshore migration. This is
              evident in the Middle Atlantic and South Atlantic bights where the difference in
              seasonal temperatures in estuaries is great. In the former, most individuals leave the
              estuary for the winter; in the latter, some portion of the population overwinters in the
              estuary. This pattern may vary with the severity of the winter.

              Naturally varying temperatures may also influence survival during the first year and
              thus impact subsequent year-class strength. Low temperatures (2 - 30C) cause
              mortality in transforming larvae and small juveniles. As a result, long and cold winters
              may cause increased mortality, especially in the northern portion of the range. Low
              temperatures also cause reduced growth and potentially increased predation rates
              because transforming larvae and juveniles may remain at smaller, more vulnerable
              sizes for longer periods.

              Dissolved oxygen influences habitat-use patterns during episodes of hypoxia     < 3
              ppm) or anoxia (0 ppm). During the 1976 hypoxia/anoxia event off New Jersey,
              migration away from this stress caused fish to concentrate in nearby estuaries and
              resulted in increased fishing mortality. If migration is not possible, natural mortality
              can result with subsequent loss of a significant portion of the local population.

              It is clear that estuaries are critical nursery areas throughout the range. High salinity,
              subtidal salt marsh creeks are the most important estuarine habitats because they








               62
                                                                                          Implications


               provide optimal conditions for growth of juveniles, especially during the spring and
               early summer of the first year. Natural or anthropogenic impacts on these creeks
               (ditching, bulkheading, etc.) probably reduce the quantity and quality of summer
               flounder nursery habitat and eventually reduce the growth and survival 'of populations
               anywhere this occurs. The same may be said@ of high-salinity- bays because these
               habitats are also used by older,juveniles and adults during the summer growing
               season.


               In the future, habitat research should focus on the transforming larvae and early
               juveniles because there is increasing evidence that year-class strength may be
               determined at this time. In addition, the fact that summer flounder are estuarine-
               dependent is critical because estuaries are the most likely to be affected by present
               and future human activities. This is especially relevant given the. increasing human
               Population levels expected on estuarine shores throughout the range of summer
               flounder.

               Resource managers who can influen,ce habitat decisions should begin to recognize
               that habitat loss and degradation has as an important effect on fisheries as
               oVerfishing, which has received disproportionately more attention. In addition, the
               habitat problem affects all fisheries, both commercial and. recreational. Further,
               because of the highly migratory nature of summer flounder populations, habitat loss
               and degradation in one local area can easily influence fish abundance elsewhere.
               Lastly, habitat research and management for summer flounder can not operate in a
               vacuum and requires complementary information on      stocks, life history, migrations,
               food habits, etc., as well as a broader understanding of habitat changes on
               geographical and decadal scales.








                Glossary

                Adult: Fish that is fully developed in morphology and meristic characteristics and has also attained
                sexual maturity.

                Allometric: Growth of a part of an organism in relation to the growth of the whole organism or some
                other part of it.

                Ambient temperature: The natural temperature of the water.

                Anoxia: Absence of dissolved oxygen in water; 0 ppm.

                Anthropogenic effects: Human impacts on the natural environment and wildlife.

                Assimilation efficiency: The rate at which an animal converts food into body weight.

                Benthic: On or pertaining to the bottom substrate (the benthos) in an aquatic habitat.

                Buoyant: Tendency to float or rise in water.

                C (OC): Temperature in degrees Celsius. To convert to Fahrenheit, use equation OF = (9/5 x OC) + 32.

                Clay: Sediments with grain size < 0.004 mm in diameter based on the Wentworth scale.

                Cohort: All the individuals resulting from the same spawning event in a population.

                Continental shelf: The edge of the continent that is submerged in relatively shallow ocean water.

                Creel survey: A survey of the recreational fishery that quantifies the fish landings at public piers and
                docks. A "creel" is the traditional satchel in which anglers pack their catch during fishing, but a creel
                survey is not restricted to fish packed in creels.

                Crustacean: Aquatic arthropods that typically have a body covered with a hard shelf, including
                lobsters, shrimps, crabs, etc.

                Diel: Occurring on a daily basis. For instance, patterns of faunal movement that are correlated with
                the cycles of light and dark are diel patterns.

                Dissolved oxygen: Oxygen that is in solution with water, and is available for plant and animal
                respiration.

                Ebbtide: Tidal stage at which ocean water flows in an offshore direction and estuarine waters flow
                toward the ocean.









                 64
                                                                                                            Glossa[y


                 Egg: A reproductive cell produced by a female organism; an ovum. May refer to both fertilized and
                 unfertilized state.


                 Efectrophoresis: A means of detecting genetic differences among populations of a species by
                 sampling proteins from members of the populations and analyzing the movement of these protein
                 particles in a medium through which an electrical charge is passed.

                 Entrainment: The intake of relatively immobile, free-floating organisms with water drawn into an
                 industrial, municipal or electric utility power plant.

                 Estuary (estuarine): Transitional environments between fresh water and salt water.

                 Etropus microstomus: Scientific name for the flatfish with the common name smallmouth flounder.
                 This is a letteye flounder in the same family (Bothidae) as summer flounder.

                 Fauna: All of the animal life in a given region or period of time.

                 Flora: All of the plant lite in a given region or period of time.

                 Flood tide: The tidal stage at which ocean water flows in an inshore direction and into estuarine
                 systems.

                 Family: In scientific classifications, an assemblage of genera possessing certain characters in common
                 by which they are distinguished. Subordinate to phylum, class and order and superordinate to genus
                 and species.

                 Genus: In scientific classifications, an assemblage of species possessing certain characters in
                 common, by which they are distinguished (plural = genera). Subordinate to phylum, class, order and
                 family and superordinate to species.

                 Gonad (gonadal): A reproductive organ or sex gland (ovary, testis) in which the gametes (ova, sperm)
                 are produced.

                 Georges Bank: A large, shallow (< 5 m imsome areas) bank on the continental shelf in the northern
                 Middle Atlantic Bight that is an important fishery region (see Fig..2.1).

                 Groundfish: Fishes that primarily inhabit the benthic environment, such as flatfish.

                 Habitat: The native environment or usual dwelling place of an animal or group of animals.

                 High tide: The period during which water depth is highest during a given tidal cycle. This occurs as
                 the terminus of flood tide, prior to the beginning of ebb tide.

                 Hypoxia: Low levels of dissolved oxygen in water       3 ppm) that are extremely stressful to most
                 aquatic life.

                 Ichthyoplankton: Very small fishes that drift in the water column as plankton and are typically larvae.






                Glossaa
                                                                                                                   65



                Inshore: Refers primarily to estuaries behind the shoreline of the coast, however may also refer to
                continental shelf areas close to the shore. Inshore movement is toward the shoreline and/or beyond
                into estuaries.


                Intertidal: Shallow areas along the shore and in estuaries that are alternately exposed and covered by
                the tides.


                Juncus: Genus name of upper estuarine plants that are generally named rushes.

                Juvenile: Young fish after attaining full adult morphology and meristic characteristics, but before
                sexual maturation.


                Larva: Young fish between time of hatching and the juvenile stage. Includes period of yolk sac
                absorption and morphological and physiological transformation.

                Low tide: The period during which water depth is lowest during a given tidal cycle. This occurs as the
                terminus of. the ebb tide, prior to the beginning of the. flood tide.

                MARMAP: Marine Resources Monitoring, Assessment and Prediction. National Marine Fisheries
                Service long-term offshore plankton survey.

                Mean low water (MLW): The average of all the low tide depths for a given region relative to a. set
                datum (reference point) in the region.

                Melanophore: Black and brown pigmented cells (chromatophores) in the epidermis (skin) of a fish that
                are capable of changing size, shape and color by expansion and contraction.

                Meristic: Numerical characteristics of the skeleton and musculature of an animal that can be used to
                identify species or races within species, such as the number of caudal (tail) fin rays.

                Middle Atlantic Bight: Zoogeographic marine region of the Atlantic Ocean that includes the estuarine
                and the continental shelf waters between Cape Cod, Massachusetts (including the northern extension
                of Georges Bank) and Cape Hatteras, North Carolina (see Fig. 2.1).

                Migration: 1. Movement of fauna from one region to another. Usually refers to regularly observed
                patterns of movement based on season and/or life history stage of a migratory animal, however, the
                term may also refer to a localized, anomalous movement as well. 2. Movement of a subset within a
                larger system, e.g., eye migration of transforming summer flounder larvae,entails the movement of the
                right eye to the left side of the head.

                Morphology (morphological): The form and structure of an organism considered as a whole.

                Morphometric: The size relationships of various morphological characteristics of an animal. For
                instance, the width vs. the depth, or the snout to eye length vs. the snout to caudal (tail) length. These
                relationships can be used to identify species or races within species.

                Mud: Sediments with grain sizes within both the clay and silt ranges of < 0.0063 mm in diameter.

                NEFSC: Northeast Fisheries Science Center.









              66
                                                                                          Glossa!y


              NMFS: National Marine Fisheries Service.


              NOAA: National Oceanic and Atmospheric Administration.

              Offshore: Refers largely to the expanse of submerged continental, shelf on the ocean side of the
              shoreline but may also. refer to areas nearer the ocean edge of the shelf. Offshore movement is away
              from the shoreline.


              Osmoregulation: The'biological strategy employed by fish to regulate and maintain their internal water
              and salt concentrations against the water and salt concentrations in their surrounding environment; i.e.,
              saltwater or freshwater.


              Overwinter: To spend the winter, Usually in a particular location or state of physiology.

              Parallchthys dentatus: Scientific name for the fish with the common name summer flounder in the
              family Bothidae (lefteye flounders).

              Pectoral fin: Paired fins located behind the head of a fish that articulate with the pectoral girdle
              (musculature).

              Pelagic: In or pertaining to the water column as distinct from the benthic region.

              Pigmentation: Coloration of tissues or cells.

              Planktonic: Pertaining to the aggregate of passively floating or drifting organisms in a body of water.

              Plankton bloom: An event where conditions are conducive to rapid reproduction of planktonic
              organisms in the ocean or other body of water.

              Population: A group of individuals of any one species that are capable of interbreeding. Uses of the
              term vary from including all individuals throughout the range to including only those individuals in a
              specified region.

              Ppm: Parts per million; units used to measure dissolved oxygen in water.

              Ppt: Parts per thousand; units to measure.the salinity of water..

              Recruitment: Addition of new individuals to a life history stage by growth and survival (e.g., from
              ,larvae to juveniles), to a specified region by movement (e.g., from offshore to estuaries), or to a fishery
              by survival or based on capture gear (e.g., mesh size).

              Ruppia maritima: Scientific name for the plant with the, common name widgeon grass. Species of
              submerged angiosperm common to estuaries in the western Atlantic. A thin, flat-bladed grass that is
              anchored to the bottom by a root-rhizome system and produces stalk-like reproductive bodies.

              Salinity: Salt content of water. The combined weight of certain salts dissolved in 1, kg of sea water,
              usually expressed as parts of salt per thousand parts of water (ppt).






               Glossaa
                                                                                                                   67



               South Atlantic Bight: Zoogeographic marine region of the Atlantic Ocean that includes the estuarine
               and continental shelf waters between Cape Hatteras, North Carolina and the southern tip of Florida
               (see Fig. 2.1).

               Scophthaimus aquosus: Scientific name for the fish with the common name windowpane flounder, a
               lefteye flounder in the same family (Bothidae) as summer flounder.

               Sand: Sediments with grain size ranging from 4.0 - @.O mm diameter (very coarse) to 0.062-0.125 mm
               in diameter (very fine) based on the Wentworth scale.

               Sift: Sediments with grain size 0.004 - 0.062 mm in diameter based on the Wentworth scale.

               Spawning: Fish reproduction process characterized by females and males depositing eggs and sperm
               into the water simultaneously or in succession so as to fertilize the eggs.

               Spartina: The genus name of salt marsh grasses generally named cordgrasses. These are the
               common tall- and short-form grasses fringing the higher-salinity portions of sandy estuarine systems of
               the western Atlantic with the Middle Atlantic being their primary range.

               Species: In scientific classifications, a group of individuals that share essential features, interbreed and
               produce similar progeny and possess the same scientific name. This name is typically two parts,
               consisting of the genus of that group (e.g., Paralichthys followed by the "specific" name (e.g.,
               dentatus . This name is unique among all life within that kingdom and is subordinate to phylum, class,
               order, family and genus.

               Stellate: Descriptive term for melanophores that appear roughly star-shaped when expanded.

               Stock: 1. A separate breeding population of a species. 2. Term used to identify a management unit of
               fishery species.

               Standard length (SL): A method of measuring fish length from the anterior part of the head to the
               posterior margin of the hypural plates.

               Stratified sampling: A study design whereby replicate sampling is conducted in predetermined
               subsets of a region, e.g., conducting the same standardized trawling program at surface, midwater and
               bottom in the water column, or at the same depth but within equal-sized blocks of an area.

               Subestuary: Smaller system within a larger estuary such as a branching subtidal marsh creek with
               interlidal tributaries.


               Substrate: General term for any benthic habitat.

               Subtidal: The shallow water zone, often only a few feet deep, which  is influenced by tides but never
               completely drained at low tide.

               Thermocline: A steep, vertical temperature gradient resulting in stratification of the water column.

               Tidal stream transport: A mechanism by which some marin      'el life passively move into estuarine
               systems by entering the water column during flood tides and moving to the bottom during ebb tides,
               resulting in net up-estuary movement.









                  68
                                                                                                             Glossa!y


                  Total length (TL): A method of measuring fish length from the most anterior part of the head to the
                  furthest extension of the caudal (tail) fin.

                  Transformation: Period of transition from the larval stage to the juvenile form characterized by both
                  morphological and physiological changes.

                  Trophic: Feeding or feeding relationships between animals.

                  Ultrasonic tag: A fish-marking tag that emits a distinct radio frequency, allowing researchers to track
                  fish movements with a receiver.


                  Ulva lactuca: Scientific name for the macroalga with the common name sea lettuce. Common to
                  estuaries, it grows in broad, translucent, bright green sheets, that are lobed or ruffled at the edges.
                  Initially attaches to the substrate with a short stalk and later drifts free.

                  Year-class: The fish spawned and hatched in a given year. Distinct from cohort in that some species
                  may have several spawning events in a given year resulting in several cohorts but one year-class.

                  Yolk sac: A bag-like ventral extension of the gut containing nutritive materials that first appears in the
                  fish embryo and is later absorbed by the larva during the stage after hatching and before feeding.

                  Young -of-the-Year: Age-O fish, or those animals born within.the past year, which have not yet
                  reached one year of age.

                  Zostera marina: The scientific name for the plant with the common name eelgrass. Species of
                  submerged angiosperm common to estuaries in the western Atlantic. A broad, flat-bladed grass that is
                  anchored to the bottom by a root-rhizome system and produces stalk-like reproductive bodies.








              'Appendix A
               Comprehensive Summer Flounder Bibliography


               1.    Abbe, G.R. 1967. An evaluation of the distribution of fish populations of the Delaware River
                     estuary. M.S. Thesis. University of Delaware. 64 pp.

               2.    Able, K.W., R.E. Matheson, W.W. Morse, M.P. Fahay and G.P. Shepherd. 1990. Patterns of
                     summer flounder (Paralichthys dentatus early life history in the Mid-Atlantic Bight and New
                     Jersey estuaries. Fishery Bulletin, U.S. 88(l): 1-12.

               3.    Adams, S.M. 1976a.   The ecology of eelgrass, Zostera marina (L.), fish communities. 1. Structural
                     analysis. Journal of Experimental Marine Biology and Ecology 22: 269-2291.

               4.             1976b. The ecology of eelgrass, Zostera marina (L.), fish communities. 11. Functional
                     analysis. Journal of Experimental Marine Biology and Ecology 22: 293-311.

               5.    Agnello, R.J. 1989. The economic value of fishing success: an application of socioeconomic
                     survey data. Fishery Bulletin, U.S. 87(l): 223-232.

               6.    AhIstrom, E.H., K. Amoaka, D.A. Hensley, H.G. Moser and BY. Sumida.    1984.
                     Pleuronectiformes: Development. Special Publication 1. Pages 640-670 in H.G. Moser, W.J.
                     Richards, D.M. Cohen, M.P. Fahay, A.W. Kendall, Jr. and S.L. Richardson, editors. Ontogeny and
                     Systematics of Fishes. American Society of Ichthyologists and Herpetologists.

               7.    Allen, D.M. and D.L. Barker. 1990. Interannual variations in larval fish recruitment to estuarine
                     epibenthic habitats. Marine Ecology Progress Series 65: 113-125.

               8.    Allen, D.M., J.P. Clymer III and S.S. Herman. 1978. Fishes of the Hereford Inlet estuary,
                     southern New Jersey. Lehigh University and The Wetlands Institute. 138 pp.

               9.    Allen, D.M., W.K. Michener and S.E. Stancyk. 1984. Pollution ecology of Winyah Bay, S.C.:
                     Characterization of the estuary and potential impacts of energy development. Baruch Institute
                     Special Publication No. 84-1. University of South Carolina, Columbia. 271 pp.

               10.   Allen, D.M., S.E. Stancyk and W.K. Michener, editors. 1982. Ecology of Winyah Bay, S.C. and
                     potential impacts of energy development. Baruch Institute Special Publication No. 82-1.
                     University of South Carolina, Columbia. 275 pp.

               11.   Almeida, F.P., R.E. Castaneda, R.V. Jesien, R.C. Greenfield and J.M. Burnett. 1992. Proceedings
                     of the NEFC/ASMFC summer flounder, Paralichthys dentatus aging workshop; 11-13 June 1990,
                     Northeast Fisheries Center, Woods Hole, Massachusetts. NOAA Technical Memorandum
                     NMFS-F/NEC-89. National Marine Fisheries Service.








                A-2
                                                                                                  Bibliography


                12.   Alperin, I.M. and J.C. Poole. 1956. Long Island's fluke. New York State Conservationist 11(3):
                      16-17.


                13.   Ames, W.H. 1954. Biological survey of the Delaware River estuary. Biennial Report 1953-1954,
                      2. University of Delaware, Marine Laboratory. pp. 21-31.

                14.   Anderson, E.D., J.M. Maso.n, A.M.T. Lange and C.J. Byrne. 1983. Cod-end mesh selectivity in
                      the Long Island spring trawl fishery for summer flounder and associated species. Document No.
                      83-33. National Marine Fisheries Service, Woods Hole Laboratory, Massachusetts. 65 pp.

                15.   Anderson, V.T., Jr. 1978. Reverse summer flounder (Paralichthys dentatus from the Middle
                      Atlantic Bight. Bulletin of the New Jersey Academy of Science 23(t): 39-41.

                16.   Anderson, W.W. 1968. Fishes taken during shrimp trawling along the South Atlantic coast of the
                      United States, 1931-35. Special Scientif ic Report-Fisheries 570. United States Fish and Wildlife
                      Service. 60,pp.

                17.   Anderson, W.W. and J.W. Gehringer. 1965. Bio log ical-stat istical census of the species entering
                      fisheries in the Cape Canaveral area. Special Scientific Report-Fisheries No. 514. United States
                      Fish and Wildlife Service. 79 pp.

                18.   Arnold, E.L. 1951. Northward dispersal of warm water fishes in southern New England during the
                      summer of 1949. Copeia 1951.

                19.   Arve, J. 1960. Preliminary report on attracting fish by oyster-shell plantings in Chincoteague
                      Bay, Maryland. Chesapeake Science 1: 58-65.

                20.   Atlantic States Marine Fisheries Commission Subcommittee for Fisheries of the Atlantic Bight.
                      1966. A summary of current research on the summer flounder. 2 pp.

                21.   Atlantic States Marine Fisheries Commission Summer Flounder Subcommittee. 1970. Life history
                      of the summer flounder. A report to the Advisory Committee. 10 pp.

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                644.  White, D.B. and R.R. Stickney. 1973. A manual of flatfish rearing. NOAA/NMFS Technical
                      Report 73-7. National Marine Fisheries Service. 42 pp.

                645.  --. 1973. A bibliography of flatfish (Pleuronectiformes) research with partial annotation.
                      Technical Report Series No. 73-6. Georgia Marine Science, Center, Skidaway Island. 76 pp.

                646.  White, J,C., Jr. and D.E. Hoss. 1964. Another record of incomplete ambicolo ratio n'in the summer
                      flounder Paralichthys dentatus. Chesapeake Science 5(3): 151-152.









                    A-48
                                                                                                                    Bibliography


                    647.   Widerstrom, F.L., Jr. 1959. An economic and financial study of the fluke otter-trawl fishery of
                           New Jersey. Commercial Fisheries Review 21: 17-26.

                    648.   Wilk, S.J. 1980. The recreational fishery. Underwater Naturalist 12(4): 40-45.

                    649.   Wilk, S.J. and B.M. Baker. 1989. Results of a fish-megainverlebrate survey of the New York
                           Bight Apex, late summer 1983. Bulletin of the New Jersey Academy of Sciences 34(2): 1-13.

                    650.   Wilk, S.J. and B.E. Brown. 1980. A description of those fisheries, which take place in the western
                           North Atlantic between the U.S.-Canadian border and North Carolina, that presently have or
                           potentially could have user group allocation conflicts. Pages 502-518 in J.H. Grover, editor.
                           Allocation of fishery resources. Proceedings of the technical consultation on allocation of fishery
                           resources, April 20-23, Vichy, France, 1980. United Nations Food and Agriculture Organization
                           and the American Fisheries Society.

                    651.   Wilk, S.J. and W.W. Morse. 1979. Annual cycle of gonad-somatic indices as indicators of
                           spawning times for 15 species of fish collected from the New York Bight. Laboratory Reference
                           SHL 79-11. National Marine Fisheries Service, Sandy Hook Laboratory, New Jersey. 54 pp.

                    652.   Wilk, S.J., W.W. Morse and D.E. Ralph. 1978. Length-weight relationships of fishes collected in
                           the New York Bight. Bulletin of the New Jersey Academy of Sciences 23(2): 58-64.

                    653.   Wilk, S.J., W.W. Morse, D.E. Ralph and T.R. Azarovitz. 1977. Fishes and associated
                           environmental data collected in the New York Bight, June 1974-June 1975. NOAA Technical
                           Report, NMFS SSRF-716. National Marine Fisheries Service.

                    654.   Wilk, S.J., W.W. Morse, D.E. Ralph and E.J. Steady. 1975. Life history aspects of New York
                           Bight finfishes. Laboratory Reference SHL 75-1. National Marine Fisheries Service, Sandy Hook
                           Laboratory, New Jersey. 265 pp.

                    655.             1976. Life history aspects of Middle Atlantic Bight finfishes. Laboratory Reference 76-3.
                           National Marine Fisheries Service, Sandy Hook Laboratory, New Jersey. 149 pp.

                    656.   Wilk, S.J., W.W. Morse and 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.
                           Fishery Bulletin, U.S. 88: 775-786.

                    657.   Wilk, S.J., R.A. Pikanowski, A.L. Pacheco, D.G. McMillan, B.A. Phelan and L.L. Stehlik. 1992.
                           Fish and megainvertebrates collected in the New York Bight apex during the 12-mile dumpsfte
                           recovery study, July 1986 - September 1989. NOAA Technical Memorandum NMFS-F/NEC-90.
                           National Marine Fisheries Service.

                    658.   Wilk, S.J. and M.J. Silverman. 1976a. Fish and hydrographic collections made by the research
                           vessels Dolphin and Delaware 11 during 1968-72 from New York to Florida. Special Scientific
                           Report-Fisheries No. 697. National Marine Fisheries Service. 159 pp.

                    659.             1976b. Summer benthic fish fauna of Sandy Hook Bay, New Jersey. NOAA Technical
                           Report NMFS SSRF-698. National Marine Fisheries Service, Sandy Hook Laboratory, New
                           Jersey. 16 pp.






                 Bibliogrgphy
                                                                                                                         A-49



                 660.   Wilk, S.J., W.G. Smith, D.E. Ralph and J. Sibunka. 1980. Population structure of summer
                        flounder between New York and Florida based on linear discrimination analysis. Transactions of
                        the American Fisheries Society 109(3): 265-271.

                 661.   Williams, A.B. 1972. A ten-year study of meroplankton in North Carolina estuaries: mysid
                        shrimps. Chesapeake Science 13(4): 254-262.

                 662.   Williams, A.B. and E.E, Deubler, Jr. 1968a. A ten year study of meroplankton in North Carolina
                        estuaries: assessment of environmental factors and sampling success among Bothid flounders
                        and Penaeid shrimps. Chesapeake Science 9(1): 27-41.

                 663.   --. 1968b. Studies on macroplanktonic crustaceans and ichthyoplankton of the Pamlico
                        Sound complex. Special Scientific Report No. 13. North Carolina Department of Conservation
                        and Development. 91 pp.

                 664.   Witting, D.A. and K.W. Able. 1993. Effects of body size on probability of predation for juvenile
                        summer flounder and winter flounder based on laboratory experiments. Fishery Bulletin, U.S. 91:
                        577-581.


                 665.   Wolff, M. 1974. Preliminary stock assessment, North Carolina flounder (Paralichthys sp.).
                        Completion Report. North Carolina Division of Marine Fisheries, Morehead City. 24 pp.

                 666.   -. 1977. Preliminary stock assessment, North Carolina flounder, (Paralichthys sp.). North
                        Carolina Division of Marine Fisheries, Morehead City. 26 pp.

                 667.   ---. 1978. Preli      'minary stock assessment, North Carolina flounder (Paralichthys sp.).
                        Completion report for Project 2-294-R. North Carolina Division of Marine Fisheries, Morehead
                        City. 20 pp.

                 668.   Woodward, A.G. 1989. Effects of mesh-size on the composition and quantity of white shrimp
                        and finfish caught with the cast net in Georgia's estuarine waters. Contribution Series No. 44.
                        Georgia Department of Natural Resources, Coastal Resources Division. 37 pp.

                 669.   Woolcott, W.S. and C. Beirne. 1967. A comparison of the skulls of young flounders in three
                        species of flounders, Paralichthys. ASB Bulletin 14(2): 46.

                 670.   Woolcott, W.S., C. Beirne and W.M. Hall, Jr. 1968. Descriptive and comparative osteology of the
                        young of three species on flounders, genus Paralichthys Chesapeake Science 9(2): 109-120.

                 671.   Wyanski, D.M. 1990. Patterns of habitat utilization in O-age summer flounder (Paralichthys
                        dentatu . M.A. Thesis. College of William and Mary, Gloucester Point. 54 pp.

                 672.   Young, B.H., K.A. McKown, V.J. Vecchio, K. Hattala and J.D. Sicluna. 1990-1991. A study of the
                        striped bass in the marine district of New York VI. Grant # NA88EA-D-AN030 (3 Reports:
                        Annual Report for Period April 1, 1988 - March 31, 1989; Annual Report for Period April 1, 1989 -
                        March 31, 1990; Completion Report for Period April 1, 1988 - March 31, 1991. New York State
                        Department of Environmental Conservation, Stony Brook.

                 673. Younger, R.R. and J.A. Zamos. 1955. New Jersey's marine sport fishery. Miscellaneous Report
                        16, Job Completion Report for Project F-2. New Jersey Division of Fish and Game.








                    A-50
                                                                                                                  Bibliography


                   671.   Zingmark, R.G. 1978. An annotated checklist of the biota of the coastal zone of South Carolina.
                          University of South Carolina, Columbia.

                   672.   Ziskowski, J.J., L.M. Despres-Patanjo, A.B. Howe, D. Ralph and S. Atran. 1987. Disease in
                          commercially valuable fish stocks in the Northwest Atlantic. Marine F@ollUtion Bulletin 18(9): 496-
                          504.









                 Author index to the Bibliography

                 Note: Numbers correspond to citations listed in Appendix A

                 Abbe, G. R., I                                              Berg, D.L., 36
                 Able, K.W., 2, 282, 293, 284, 516, 517, 518, 603,           Berg, M.B., 37
                       604,605,664                                           Berrien, P.L., 97
                 Adams, S.M., 3, 4, 608                                      Berry, F.H., 635, 636
                 Agnello, R.J., 5                                            Sevacqua, E.S., 23
                 AhIstrom, E.H., 6,224                                       Bieder, R.C., 38
                 Allen, D.M., 7, 8, 9, 10, 449                               Bigelow, H.B., 39
                 Almeida, F.P., I I                                          Birdsong, R.S., 107
                 Alperin, I.M., 12                                           Bisbal, G.A., 40
                 Ames, W.H., 13                                              Bodammer, J.E., 41, 311
                 Amoaka, K., 6                                               Boehlert, G.W., 452
                 Anderson, E.D.,    14                                       Bolz, G.R., 42
                 Anderson, V.T. Jr.,   15                                    Bond, C.E., 503, 504
                 Anderson, W.W., 16, 17                                      Sonzek, C.F., 43, 44, 190
                 Angelovic, J.W., 471                                        Bowman, E., 209
                 Arnold, E.L., 18                                            Bowman, R.E., 151, 302, 303, 347
                 Arve, J., 19                                                Bozeman, E.L. Jr., 45
                 Atlantic States Marine Fisheries Commission                 Breder, C.M. Jr., 46, 47, 436
                 Subcommittee for Fisheries of                               Briggs, J.C., 48
                       the Atlantic Bight, 20                                Briggs, P.T., 49, 50, 51, 52
                 Atlantic States Marine Fisheries Commission                 Brooker, JR, 503, 504
                 Summer Flounder                                             Brooks, H.A., 630
                       Subcommittee, 21, 22                                  Brown, BE, 88, 89, 650
                 Atran, S., 672                                              Brown, C., 506, 507
                 Ayvazian, S.G., 126                                         Brown, R.A., 53
                 Azarovitz, T.R., 23, 24, 74, 208, 653                       Brown, J.T., 220
                 Bailey, R.M., 26, 503, 504                                  Bruce, R.A., 54
                 Baker, B.M., 649                                            Buchanan, C.C., 55,461
                 Baker, S., 147                                              Buckley, L.J., 56, 57
                 Baptist, J.P., 27                                           Buller, R.J., 58
                 Barans, C.A., 635, 636                                      Bullis, H.R., 59
                 Barker, D.L., 7                                             Bulloch, D.K., 60
                 Bass, D.G. Jr., 28                                          Burke, J.S., 61, 62, 63', 376, 624
                 Bayless, J., 291                                            Burnett, J.M., 11, 446
                 Bean, B.A., 611                                             Burns, R.W., 64
                 Bean, T.H., 29                                              Burreson, E.M., 65, 66, 67, 68, 69, 70, 71, 72,
                 Bearden, C.M.,   30t 3 1, 32, 33                                  263,602
                 Beatty, H.R., 34, 637                                       Bushing, M.F., 73
                 Bedsole, H.L. Jr., 35                                       Byrne, C.J., 14, 23, 24, 74
                 Beirne, C., 669, 670                                        Byron, R.R., 96
                 Bengston, D.A., 40                                          Cable, L.E., 231, 232








                     A-52
                                                                                                                            Bibliography


                     Cable, R.M., 75                                                   Desfosse, J.C., 134, 125
                     Cain, R.E., 76                                                    Despres, L.I., 23
                     Cali, A., 306                                                     Despres-Patanjo, L.I., 24, 135, 136, 672
                     Carley, D.H., 78                                                  de Sylva, D.P., 137, 138, 545
                     Carlson, J.K., 79                                                 Deubler, E.E. Jr., 140, 141, 662, 663
                     Carolina Power and Light Co., 80                                  Deuel, D.G., 142, 143, 323
                     Casey, J.F., 81                                                   DeVries, D.A., 144, 145, 513
                     Castagna, M., 501                                                 Dickinson, C.L., 427
                     Castaneda, R.E., I I                                              Dillmann, DA, 57
                     Chadwick, D.L., 108, 110                                          Dipiero, S.J., 564
                     Chang, S., 82, 83                                                 Donohoe, C.J., 199
                     Chester, A.J., 229                                                Douglas, J., 505
                     Christensen, D.J.,      84, 85, 92, 185                           Dovel, W.L., 146
                     Clark, J.R., 86, 87, 143                                          Draxler, S., 593
                     Clark, S.H., 88, 89                                               Drewry, G.E., 342
                     Clayton, G., 90                                                   Dudley, D.L., 606
                     Clements, L.C., 91                                                Duncan, J., 540, 541
                     Clifford, W.J., 84,85,92                                          DuPaul, W., 147
                     Clymer, J.P. 111, 8                                               Easley, J.E. Jr., 106
                     Coates, P.G., 172                                                 Ecological Analysts Engineering Science and
                     Coduri, R.J., 93                                                  Technology, Inc., 148
                     Cole, C., 90                                                      Ecological Analysts, Inc., 149
                     Cole, R.W., 94,95,                                                Edwards, R.L., 150, 151, 152
                     Colton, J.B. Jr., 96,   97,98,334,335,336                         Eerly, S.P., 153, 154, 515
                     Colton, M.D., 99                                                  Eisenberg, M., 159
                     Colvocoresses, J.A., 43, 44, 100, 190                             Eklund, A., 155
                     Connecticut Division of Marine Fisheries,         101,            Eldridge, P.J., 156
                            102,103                                                    Essig, R.J., 157
                     Conover, D.O., 363                                                Estes, A.D., 134, 135
                     Cooke, D.W., 199                                                  Erickson, M.C., 125
                     Coon, W.P. 11, 639, 640                                           Estes, A.D., 406
                     Copeland, B.J., 104, 105, 227                                     Estrella, B.T., 249
                     Coston, L.C., 246, 247                                            Evermann, B.W., 275
                     Coston-Clements, L.C., 425, 530, 531                              Fahay, M.P., 2, 87, 97, 158, 386, 387, 580
                     Cowan, J.H. Jr., 107                                              Fahy, W.E., 140
                     Curley, J.R., 108, 109, 110, 171                                  Farmanfarmaian, A., 159
                     Dahlberg, M.D., if 1, 112, 113, 114, 115,                         Farmer, C.H. 111, 32
                     Daiber, F.C., 116, 117, 118, 119, 120, 573                        Fee, R., 160
                     Daniel, L.B. 111, 121, 638                                        Feigenbaum, D.L., 73
                     Davidson, L., 540, 541                                            Festa, P.J., 161, 162, 163, 164, 165, 166, 167,
                     Davis, R.W., 631                                                          168,396,397,398
                     Dawson, C.E., 122, 123                                            Figley, W., 169, 170
                     Deacutis, C.F., 124                                               Finn, J.T., 126
                     Dean, J.M., 45, 77, 553                                           Fiske, J.D., 109, 171, 172
                     Decker, E.A., 125                                                 Fitch, J.E., 26
                     Deegan, L.A., 126                                                 Fitzhugh, G.R., 376
                     DeGroot, S.J., 127,     128, 129                                  Florida Department of Natural Resources, 173
                     Delaney, G.R., 130,     175                                       Fogarty, M.J., 174, 175
                     Derickson, W.K., 131                                              Foster, D.B., 335, 336
                     Dery, L.M., 132, 133, 466, 574                                    Foster, H.A., 23, 24





                   QibliograRhy                                                                                                 A-53


                   Fowler, H.W., 176, 177, 178, 179, 180, 181, 182,               Hattala, K., 672
                          183, 551, 611                                           Hawkes, JA, 239
                   Freeman, B.L., 184, 185, 186                                   Hawkins, J.H., 221, 513
                   Frisbie, C.M., 78                                              Heck, K.L. Jr., 453
                   Frizzell, L.J., 68                                             Henderson, E.M., 22
                   Gabriel, W.L., 187                                             Hennemuth, R.C., 223
                   Gahn, T., 594, 595, 596, 598                                   Hensley, D.A., 6, 224
                   Gaichas, S.K., 270                                             Herald, E.A., 26
                   Gallagher, M.L., 188                                           Herman, S.S., 8, 225, 286
                   Gartner, J.V., 189                                             Herold, R.C., 226
                   Geer, P.J., 43, 44, 190                                        Herrema, D.J., 199
                   Gehringer, J.W., 17                                            Hester, J.J., 227
                   Gerhart, E.H., 191                                             Hettler, W.F. Jr., 228, 229, 247, 609
                   Gerry, L.R., 632                                               Heyerdahl, E.G., 210
                   Gibson, C.I., 355                                              Hickey, J.M., 108, 109
                   Gilbert, C.R., 192                                             Hicks, B., 230
                   Gillikin, J.W. Jr., 35, 175, 193, 194, 195, 196, 482,          Hildebrand, S.F., 231, 232, 233
                         483                                                      Hill, N.L., 220
                   Gilmore, R.G. Jr., 197, 198, 199, 274                          Hillman, R.J., 234
                   Ginsburg, L, 200                                               Himchak, P.J., 235, 236, 237, 238
                   Ginter, J.J.C., 364                                            Hocuff, C.H., 269, 270
                   Goldman, J.C., 201                                             Hodgins, H.O., 239
                   Goldstein, R.J., 202                                           Hodson, R.G., 104, 105, 106, 632
                   Goode, G.B., 203                                               Hoese, H.D., 240
                   Gordon, B.L., 204                                              Hoff, F.H. Jr., 610
                   Goss, D.K., 276                                                Hoff, J.G., 575
                   Grant, G.C., 458,459                                           Hoffman, W.G. 11,    241
                   Graunke, B., 274                                               Hogarth, W.T., 539
                   Greenfield, R.C., I I                                          Holder, D.R., 248
                   Greenwood, P.H., 205                                           Holland, A.F., 312
                   Greges, M.P., 564                                              Holland, B.F. Jr., 35, 195, 196, 242, 482, 483
                   Grimes, B.H., 206                                              Holliday, M.C., 157
                   Grosslein, M.D., 25, 207, 208, 209, 2 10                       Horwitz, R.J., 243
                   Gudger, E.W., 211, 212, 214                                    Hoss, D.E., 62, 63, 91, 244, 245, 246, 247, 609,
                   Gunter, G., 214                                                       646
                   Gusey, W.F., 215                                               Hottell, H.E., 248
                   Gutherz, E.J., 216                                             Howe, A.B., 249, 671
                   Guthrie, R.O., 195                                             Howell, W.H., 271, 272
                   Hackney, C.T., 520                                             Hsia, R., 159
                   Haire, M.S., 312                                               Hughes, E.H., 250,251
                   Halgren, B., 217                                               Huish, M.T., 206
                   Hall, W.M, Jr., 670                                            Hultin, H.O., 125
                   Hamer, P.E., 152, 218, 314                                     Hunninen, AN., 75
                   Hardy, J.D. Jr., 333                                           Hussakoff, L., 252
                   Hargis, W.J. Jr., 219                                          lannaccone, V., 159,
                   Harrell, M.L., 188                                             Ichthyological Associates, 254, 255, 256, 257,
                   Harris, C.D., 324, 325, 326, 327                                      258
                   Harris, R.E., 43, 44, 190                                      Industrial Fishery Project, 259
                   Harvell, C.H., 145                                             International Gametish Association, 260
                   Hassler, W.W., 220                                             Irlandi, L.R., 425








                  A-54
                                                                                                                Bibliocjrap@y


                  Ishibashi, N., 261                                           Leim, A.H., 308
                  Itzkowitz, N., 262, 564                                      Lesser, C.A., 309
                  Jacobs, F., 312                                              Levinton, J.S., 36
                  Jansen, M.E., 263                                            Lewis, E.J., 310, 311
                  Jearld, A. Jr., 574                                          Lewis, R.R., 562
                  Jeffries, H.P., 264, 265                                     Lindsey, C.C., 26
                  Jensen, A.C., 266, 267, 268                                  Lipcon, V., 598
                  Jensen,J., 312                                               Lison, AA, 312, 313,
                  Jesien, R.V., 11, 269, 270                                   Livingstone, R., 152
                  Johns, D.M., 271, 272                                        Livingston, R.J., 556
                  Johnson, G.N., 612, 613, 614                                 Lockwood, K., 170
                  Johnson, K.L., 273                                           Longo,J., 564
                  Johnson, W.C., 264                                           Lough, R.G., 42
                  Jones, R.S., 274                                             Low, R., 33
                  Jordan, D.S., 275, 276                                       Lucy, J.A., 341
                  Jorgenson, S.C., 277, 375                                    Lugger, 0., 616
                  Joseph, E.B., 344, 345, 552                                  Lux, F.E., 218, 314, 315, 316, 317, 318, 319
                  Judy, M.H., 278                                              Lynch, T.R., 320
                  June, F.E., 279                                              Lyons, P., 135
                  Junqueira, L.C.,    280,281                                  MacKenzie, C.L. Jr., 321
                  Kalber, F.A., 137, 138                                       MacPhee, G., 322
                  Kasper, V., 594, 595, 596, 598                               Maddox, M.B., 638
                  Keefe, M., 282, 283, 284                                     Magley, W.C., 274
                  Keirans, W.J. Jr., 285, 286                                  Mahoney, J.B., 315, 323, 561
                  Keiser, R.K., 287                                            Mahood, R.K., 324, 325, 326, 327
                  Keisler, A.S., 626                                           Malins, D.C., 618
                  Kernmerle, S., 288                                           Malloy, K.D., 328, 329, 330, 331
                  Kendall, AW Jr., 87, 97, 289                                 Malsberger, R.G., 296
                  Kerby, J.H., 206                                             Mann, W.C., 310
                  Kernehan, R.J., 622                                          Manooch, C.S. 111, 332
                  Kerr, G.A., 290                                              Mansueti, A.J., 333
                  Keup, L., 291                                                Marak, R.R., 334, 335, 336
                  Kimmel, J.J., 292                                            Marcellus, K.L., 337
                  Kindred, J.E.,      1971                                     Marsh, E., 338
                  Kirsch, R., 294                                              Marshall, A., 339, 340, 341
                  Kielson, M.A., 472, 609                                      Martin, F.D., 342
                  Klein-MacPhee, G., 272, 295, 296, 297                        Martore, R.M., 583, 637
                  Krouse, C.W., 530, 531                                       Mason, D., 540,541
                  Kulczyck, G.R., 274                                          Mason, J.M., 14
                  Kuntz, A., 298                                               Massmann, W.H., 343, 344, 345, 490
                  Kyle, H.M., 299                                              Mast, S.O., 346
                  Lachner, E.A., 26, 503, 504                                  Matheson, R.E., 2
                  LaCroix, M.W., 608, 609                                      Maurer, R.O. Jr., 347
                  Lange, A.M.T., 14, 300                                       Mayo, R.K., 348, 349, 446, 562
                  Langton, R.W., 301, 302, 303,                                McAdams, M., 540, 541
                  Lascara, U., 304                                             McAllister, D.E., 350
                  Latham, R., 105                                              McClain, J.F. Jr., 351, 352
                  Laudan, R., 306                                              McDermott, V., 353, 354
                  Laurence, G.C.,     307                                      McEachran, J.D., 407
                  Lawton, R.P., 108, 109,   110, 171                           McErlean, A.J., 355






                   Bibliography
                                                                                                                               A-55



                   McGovern, J.C., 356, 357, 637                                  Newman, M.W., 434
                   McHugh, J.L., 358, 359, 360, 361,,362, 363, 364                Nichols, J.T., 435, 436
                   McKenzie, R.A., 620                                            Nicholson, N., 460
                   McKown, K.A., 672                                              Nichy, F.E., 316, 317, 318
                   McLaughlin, S.M., 311                                          Nixon, S.W., 456
                   McMillan, D.G., 582, 657                                       Noble, E.B., 437, 438
                   Mercer, L.P., 365                                              Norcross, J.J., 344, 345
                   Meredith, A., 288                                              Norman, J.R., 439
                   Merriman, D., 366, 623                                         North Carolina Division of Marine Fisheries,
                   Metzger, F. Jr., 367                                                  440-444
                   Michels, S.F., 368                                             Northeast Utilities Service Company, 445
                   Michener, W.K., 9, 10,449                                      O'Brien, L., 446
                   Mid-Atlantic Council Staff, 369                                O'Conner, J.S.,     447,448
                   Mid-Atlantic Fishery Management Council,         370,          O'Connor, S.G., 355
                          371,372,373,374                                         Odum, E.P., 115
                   Midlige, F.H., 323                                             Ogburn, M.V., 449
                   Miglanese, J.V., 552                                           Olla, B.L., 450
                   Mihursky, J.A., 355                                            Olne y, J.E., 451, 452
                   Miller, D., 336, 590                                           O'Neil, S.P., 575
                   Miller, G.L., 277, 375                                         Orth, R.J., 453
                   Miller, J.M., 376, 377, 378                                    Osborn, C.M., 454, 455
                   Miller, J.S., 63                                               Oviatt, C.A., 456
                   Miller, R.W., 379                                              Pacheco, A.L., 83, 457, 458, 459, 657
                   Monaco, M.E., 425                                              Pafford, J.M., 402, 460
                   Monaghan, J.P. Jr., 365, 380a,     380b,491                    Palmer, B.A., 324, 325, 326, 327_
                   Monroe, R.J., 104, 437, 438                                    Pancirov, R.J., 53
                   Moore, H.F., 381                                               Parker, L.L., 73
                   Moran, D., 206                                                 Parker, R.O., 461
                   Moran, J.E., 638                                               Parrish, J., 90
                   Moran, R.L., 313                                               Pearcy, W.G.,       462
                   Moran-Johnson, R.L., 312                                       Pearson, J.C.,      463,464
                   Morgan, M.A., 564                                              Pendleton, E.C., 465
                   Morris, T.L. Jr., 382                                          Pentilla, A., 466
                   Morse, W.W., 2, 383, 384, 385, 386, 387, 65 1,                 Perlmutter, A., 467, 468
                          652,653,654,655,656                                     Perra, P., 469
                   Moser, H.G., 6                                                 Perschbacher,       P., 540, 541
                   Murawski, S., 90, 388, 389, 390, 391, 392, 393,                Peters, D.S., 377, 470, 471, 472, 530, 531
                          394, 395, 396, 397, 398, 399                            Phalen, P.S., 473
                   Murchelano, R.A., 136, 400, 401, 507, 523                      Phelan, B.A.,       657
                   Music, J.L. Jr., 324, 325, 326, 327, 402, 403                  Pietrafesa, L.J., 378
                   Musick, J.A., 100, 134, 135, 404, 405, 406, 407                Pikanowski, R.A., 657
                   Myer, G.S., 205                                                Podgar, T.T., 312
                   Myrvik, O.N., 626                                              Poole, J.C., 12, 175, 314, 474, 475, 476, 477,
                   Nagle, J.J., 593, 594, 595, 596, 599                                   478,479
                   National Marine Fisheries Service,      408-423                Porter, H.J., 542
                   Nelson, D.M., 425                                              Porter, L.R. Jr., 319
                   Nesbit, R.A., 426                                              Potter, D.C., 42
                   Neville, W.C., 42r6, 427, 642                                  Powell, A.B., 480, 481, 482, 483, 484, 485, 486
                   New Jersey Department of Environmental,                        Powles, H., 487
                          Protection, 428-433                                     Price, K. Jr., 131







                   A-56
                                                                                                                   Bibliograghy


                   Price, K.S., 545                                              Schaefer, R.H., 532
                   Price, T.J., 27                                               Schauss, R.P., Jr., 533
                   Pritchard, E.S.,    24                                        Schroeder, W.C., 39, 233, 534
                   Pulley, M.G., 488                                             Schubel, J.R., 262, 564
                   Purvis, C., 489                                               Schwartz, F.J., 494, 485, 486, 535't 536, 537, 538,
                   Radcliffe, L., 298                                                  539,540,541,542
                   Ralph, D.E., 175,   652, 653, 654, 655, 660, 672              Sclar, R.C., 366
                   Rand, H.G. Jr., 93                                            Scott, W.B., 26, 308, 543
                   Raney, E.C., 490.                                             Scott, R.N., 503, 504
                   Ray, G.C., 505                                                Scotton, L.N., 544, 545
                   Raynie, R.C., 81                                              Seagraves, R.J., 94, 95, 546, 547, 548, 549
                   Reback, K., 108,    110                                       Sedberry, G.R., 634
                   Reed, J.P., 378                                               Settle, M.E., 425
                   Reed, L., 491                                                 Sharp, B., 551
                   Reichert, M.J.M.,   492                                       Sharp, J.H., 550
                   Reinach, F., 280                                              Shealy, M.H. Jr., 552, 639, 640, 641
                   Reintjes, J.W., 279, 493                                      Shenker, J.M., 553
                   Reis, R., 494                                                 Shepherd, G.P., 2, 85, 554, 555
                   Reisersen, L.,      495                                       Sheridan, P.F., 556
                   Rhode Island Division of Fish and Wildlife,    .496           Sherman, K., 557, 558
                   Rhodes, R., 33                                                Sherr, E.B., 251
                   Rice, C.D., 627                                               Shipman, S., 559
                   Richards, A., 497                                             Shuster, C.N. Jr., 138
                   Richards, C.E., 498, 499, 500, 501                            Shuster, C.W., 560
                   Richards, S.W.,     462,502                                   Sibunka, J.D., 581, 660
                   Richkus, W.A.,      312                                       Sicluna, J.D., 672
                   Riggs, S.R., 105, 106                                         Silverman, M.J., 561, 658, 659
                   Robins, C.R., 503, 504, 505                                   Sinderman, C.J., 601
                   Robohm, R.A., 506, 507, 508                                   Sissenwine, M.P., 562
                   Rogers, S.G., 403, 509, 5 10                                  Smigielski, A.S., 563
                   Roithmayr, C.M., 493                                          Smith, C.F.,   564
                   Rosen, D.E., 205                                              Smith, H.M.,   565,566,567
                   Rosenberg, M., 5 L i                                          Smith, J.W.,   568t569,638
                   Ross, J,L., 365, 491, 512, 513                                Smith, M.G.,   543
                   Ross, S.W., 154, 514, 515                                     Smith, N.S.,   545
                   Rournillat, W.A., 637, 638                                    Smith, R.E.,   545
                   Rountree, R.A., 516, 517, 518,   605                          Smith, R.L.,   570
                   Rozas, L.P., 519, 520                                         Smith, R.W.,   120, 175, 571, 572, 573, 574,
                   Rulifson, R.A., 188, 521, 522                                 Smith, S.I., 619
                   Ryder, C., 126                                                Smith, S.M., 575
                   Salles, L.M.M.,     280,281                                   Smith, W.G.,   87, 97, 386, 387, 558, 576-582, 66.0
                   Samet, C.E., 450                                              Socci, R., 159
                   Sandifer, P.S., 639, 640,  641                                Sparrow, D.S., 508
                   Sandoy, K., 540, 541                                          Spear, H.S., 58
                   Sass, S.L., 523                                               Spitsbergen, D., 582
                   Sawyer, R.T., 524                                             Stancyk, S.E., 9, 10
                   Sawyer, T.K., 311                                             Steady, E.J., 654, 655
                   Scarlett, P.G., 175, 217, 525, 526, 527,  528, 574,           Stehlik, L.L., 656, 657
                   Scattergood, L.W., 529                                        Stender, B.W., 583, 635, 636
                   Schaaf, W.E., 246, 530, 531                                   St. Onge, J.M., 98





                 Bibliogrgphy
                                                                                                                           A-57



                 Stender, B.W., 487                                            Weinstein, M.P., 539, 575, 628, 629, 630, 631,
                 Stephan, C.D., 584, 585                                             632, 633
                 Stephan, D.C., 473                                            Weiss, S.L., 632,633
                 Stern, H., Jr., 210                                           Weitzman, S.H., 205
                 Stickney, R.R., 586, 587, 588, 589, 590, 644, 645             Weller, P., 37
                 Stock Assessment Workshop, 591, 592                           Wells, A., 581, 582
                 Stolen, J.S., 306, 593, 594, 595, 596, 597, 598,              Wenner, C.A., 33, 357, 569, 634, 635, 636, 637,
                 Stone, R.B., 461                                                    638
                 Struhsaker, P., 599                                           Wenner, E.L., 339 34, 639, 640, 641
                 Studholme, A.L., 450                                          Wesche, A.E., 81
                 Sumida, B.Y., 6                                               Westman, J.R., 427, 642
                 Sumner, F.B., 600                                             Wheatland, S.B., 643
                 Swanson, R.L., 601                                            White, D.B., 587, 588, 589, 590, 644, 645
                 Sypek, J.P., 69, 602                                          White, J.C. Jr., 141, 646
                 Szedlmayer, S.T., 603, 604, 605                               White, R.L., 399
                 Tagatz, M.E., 606                                             Widerstrom, F.L. Jr., 647
                 Targett, T.E., 155,   329, 330, 331, 509                      Wilk, S.J., 175, 648-660
                 Tate, J., 541                                                 Williams, B.C., 403
                 Temple, R.F., 99                                              Williams, A.B., 661, 662, 663
                 Terceiro, M., 265, 555, 607                                   Witting, D.A., 664
                 Thayer, G.W., 608,609                                         Witzig, J.F., 157
                 Thompson, J.R., 59                                            Wolff, M., 665, 666, 667
                 Tinsman, J.C., 94, 95                                         Woodward, A.G., 668
                 Topp, R.W., 610                                               Woolcott W.S., 669,670
                 Truitt, R.V., 611                                             Wyanski: D.M., 671,
                 Turner,W.R., 612,613,614                                      Young, B.H., 672
                 Tyler, AN., 615                                               Younger, R.R., 673
                 Uhler, P.R., 616                                              Zamos, J.A., 673
                 Van Den Avyle, M.J., 510                                      Zingmark, R.G., 671
                 Van Der Veer, H.W., 492                                       Ziskowski, J.J., 400, 401, 672
                 Van Dolah, R., 33                                             Zwerner, D.E., 70, 71, 72
                 Van Housen, G., 617
                 Van Sant, S.B., 509
                 Varanasi, U., 618
                 Vaughn, D.S., 530, 531
                 Vecchio, V.J., 672
                 Verrill, A.E., 619
                 Vladykov, V.D., 620
                 Vouglitois, J.J., 621
                 Walford, L.A., 186
                 Walters, M.F., 633
                 Wang, J.C.S., 622
                 Warfel, H.E., 623
                 Warlen, S.M., 624
                 Warriner, J.E., 626, 627
                 Watson, C.E., 172
                 Weber, A.M., 625
                 Webster, R.P., 34
                 Weeks, B.A., 626, 627








                Subject index to the Bibliography


                Note: Numbers correspond to citations listed in Appendix A


                adults. 144
                age: 11, 132, 133, 466, 474, 481, 554
                anatomy: 128, 129, 226, 280, 281, 293, 294, 299, 338, 382, 455,
                      669, 670 (see also anomalies)
                anomalies: 15, 40, 111, 123, 140, 189, 211, 212, 213, 252, 494, 589, 646
                aquaculture: 587, 588, 589, 644
                Baltimore. Canyon: 53
                behavior: t27, 295, 296, 304, 346, 450, 590
                bibliography: 123, 495, 526, 545, 645
                biochemistry: 37, 125, 159, 188, 330, 331, 511
                biomass: 88, 319, 516, 652
                Block Island Sound: 366
                Canada: 97, 98, 158, 187,  308, 350, 384, 386, 387, 420, 543, 620, 650
                Chesapeake Bay: 67, 69, 70, 71, 72, 73, 121, 233, 243, 263, 333, 404, 452, 453, 464, 469
                Chesapeake Bight: 407, 468
                color: see pigment
                commercial fisheries: see fisheries
                Connecticut: 79, 101, 102, 103, 462
                Delaware: 131, 331, 362, 368, 379, 458, 459, 544, 546
                Delaware Bay-River: 1, 13, 1.20, 137, 138, 177, 191, 238, 254, 257, 279, 433,
                      493, 545, 547, 548, 549, 560, 570, 571, 572, 573, 622
                development: 6, 41, 205, 231, 232, 283, 298, 299, 313, 333, 342, 455
                disease/parasites: 65, 66, 67, 68, 69, 70, 71, 72, 75, 122, 136, 219, 239,
                      263, 306, 311, 323, 400, 401, 4.34, 506, 507, 508, 5239 524, 561, 588, 593, 594, 595, 596, 597, 598, 602,
                      672
                distribution (temporal & spatial): 1, 23, 24, 25, 34, 42, 48, 64, 76, 77, 98, 115, 150, 208, 425, 229, 230, 264,
                      289, 291, 334, 335, 336, 355, 396, 405, 407, 456, 462, 485, 487, 517, 521, 544, 552, 553, 562, 576, 579,
                      581, 599, 629, 634, 639, 640
                dredging: 119,534,586
                early life history: 2, 158, 376, 458, 510, 622








                   A-60
                                                                                                                    Bibliography


                   eggs: 2, 42, 96, 98, 146, 225, 229, 237, 241, 271, 272, 278, 285, 286, 298, 333, 3.14, 335, 336, 352, 354, 366,
                           451, 452, 457, 467, 487, 557, 576, 577, 580, 582, 621, 637, 643
                   electrophoresis: 93, 617
                   entrainment: see industry/energy plant.studies
                   estuarine dependence: 87, 378
                   fecundity: see reproduction
                   feeding: 61, 128, 129, 261, 292, 304, 328, 329, 330,    382, 450, 477, 590
                   fisheries (commercial/recreational): 5, 14, 16, 17, 20, 21, 22, 32, 33, 55, 58, 60, 78, 82, 84, 85, 86, 92, 94,
                           95, 116, 117, 118, 142, 143, 147, 155, 156, 157, 160, 162, 163, 164, 165, 166, 167, 168, 169, 170, 173,
                           174, 185, 186,  203, 210, 235, 249, 259, 260, 266, 267,  268,  273., 279, 287, 288, 300, 309, 316, 317, 320,
                           324, 325, 326,  327, 339,  340, 341, 351, 359, 360, 361,  361),  363, 364, 365, 369, 370, 371, 372, 373, 374,
                           379, 394, 398, 402, 403, 408, 409, 410, 411, 412, 413, 414, 415, 417, 418, 419, 421, 422, 423, 426, 438,
                           441, 442, 443, 460, 463, 468, 469, 473, 475, 476, 491, 493, 496, 498, 499, 512, 513, 514, 525, 527, 532,
                           546, 548, 549,  555, 557,  558, 559, 568, 599, 625, 642,  647,  648, 650, 672
                   Florida: 28, 48, 173, 192, 197, 198, 199, 186, 274, 290, 556, 658, 660
                   food/iood habits: 73, 129, 151, 167, 236, 250, 251, 292, 301, 302, 303, 347, 405, 470, 471, 472, 486, 619
                   Georges Bank: 42, 96, 99, 334, 335, 336, 579
                   Georgia: 78, 111, 112, 113, 114, 115, 240, 248, 251, 277, 324, 325, 326, 327, 375, 402, 403, 460, 492, 509,
                           559,668
                   Gulf of Maine: 39, 89, 96, 334, 335, 336, 529
                   growth: 56, 261, 282, 295, 296, 329, 329, 330, 331, 470, 471, 475, 491, 492, 517, 574, 603, 605
                   habitat selection/use: 61, 62, 282, 322, 437, 510, 520, 5-34, 671
                   Hudson River: 36, 146
                   identification: 47, 93, 114, 158, 200, 204, 216, 224, 233, 275, 276, 308, 332, 439, 505,,545, 610, 620
                   immunology: see physiology
                   impingement: see industry/energy plant studies
                   industry/energy plant studies: 9, 10, 80, 104, 112, 124, 148, 149, 234, 247, 254, 255, 256, 257, 258, 337,
                           354,367,564
                   juveniles (incl. post-larvae): 1, 2, 43, 44, 45, 68, 104, 107, 139, 141, 161, 190, 138, 245, 282, 328, 329, 330,
                           331, 333, 356, 357, 378, 396, 457, 458, 459, 464, 470, 471, 472, 492, 494, 533, 547, 548, 553, 577, 578,
                           602, 603, 604, 605, 609, 632, 664, 671
                   larvae: 2,  7, 41, 42, 45, 56, 62, 64, 91, 96, 98, 104, 107, 146, 158, 161, 225, 229, 237, 238, 241, 246, 247,
                           271, 272, 278, 285, 286, 289, 297, 298, 307, 310, 333, 334, 335, 336, 344, 345, 353, 354, 356, 357, 366,
                           378, 396,451,452,457,459,     467, 487, 494, 533, 544, 545, 553, 557, 563, 576, 577, 580, 581, 582, 621,
                           624, 637, 643
                   length (incl. length/weight): 92, 169, 277, 307, 317, 319, 348, 375, 552, 652
                   life history: 21, 22, 25, 38, 192, 203, 206, 231, 232, 322, 402, 442, 479, 530, 531, 621, 638, 642, 654
                   Long Island Sound: 427, 445, 502
                   marking: see tagging/marking
                   Maryland: 19, 81, 179, 269, 270, 312, 313, 535, 536, 537, 611, 616






                  Biblioargphy
                                                                                                                              A-61



                  Massachusetts: 90, 108, 109, 110, 126, 172, 249, 317, 523, 551, 566, 576, 581
                  maturity: see reproduction
                  mesh selectivity: 14, 194, 195, 217, 668
                  meristics: 130
                  metabolism: see physiology
                  metamorphosis: 283, 299
                  Middle Atlantic: 2, 23, 24, 58, 60, 65, 134, 135, 155, 175, 184, 206, 215, 289, 314, 316, 342, 360, 363, 383,
                         405, 406, 410, 468, 550, 555, 582, 615, 655
                  migration: see movement
                  morphometrics: 130
                  mortality/survival: 65, 72, 297, 328, 329, 330, 492, 601, 605, 625
                  movement: 145, 193, 218, 242, 245" 269, 270, 305, 378, 380a, 604 (see also tagging/ marking)
                  Nantucket Shoals: 42, 152, 186
                  New England: 18, 150, 266, 318, 436, 555, 623
                  New Haven Harbor: 79
                  New Jersey: 2, 8, 29, 46, 84, 85, 148, 149, 161, 162, 163, 164, 165, 166, 167, 168, 169, 170, 176, 178, 180,
                         181, 182, 183, 184, 185,  234, 235, 236, 237, 238, 255, 256, 258, 285, 286, 288, 321, 337, 351, 352, 353,
                         354,  367, 381, 388, 389, 390, 391, 392, 393, 394, 395, 396, 398, 428, 429, 430, 431, 432, 433, 516, 517,
                         518, 528, 565, 603, 604,  605, 647, 673
                  New York:     12, 14, 49, 50, 51, 52, 98, 49, 267, 321, 435, 436, 467, 474, 475, 476, 477, 478, 532, 564, 625,
                         642, 658, 660, 672, 358
                  New York Bight: 55, 74, 323,     361, 364, 400, 401, 447, 448, 507, 561, 601, 649, 651, 652, 653, 654, 656, 657
                  nitrogen:  57
                  North Atlantic: 54, 59, 83, 88,  136, 158, 174, 187, 209, 216, 222, 223, 301, 302, 303, 307, 349, 382, 411, 414,
                         419, 424, 466, 557, 558,  615, 650, 672
                  North Carolina: 35, 80, 89, 97, 104, 105, 106, 139, 141, 144, 145, 152, 153, 154, 158, 187, 193, 194, 195, 196,
                         220, 227, 228, 229, 232, 241, 242, 253, 278, 291, 310, 331, 377, 380b, 384, 386, 387, 420, 437, 438, 440,
                         441, 442, 443, 444, 463, 465, 472, 473, 480, 482, 483, 485, 486, 488, 489, 512, 513, 514, 51-5, 519, 521,
                         538, 539, 540, 541, 542, 567, 576, 581, 582, 584, 585, 606, 613, 624, 628, 650, 661, 662, 663, 665, 666,
                         667
                  nursery habitat: 61, 63, 221, 330, 458, 489, 509, 515, 628, 671
                  osteology: see anatomy
                  oyster-shell habitat: 19
                  oxygen: 91, 184, 550
                  parasites: see disease/parasites
                  physiology: 159, 280, 281, 293, 294, 295, 296, 306, 454, 455, 593, 594, 595, 596, 597, 598, 600, 602, 618,
                         626,627
                  pigment: 140, 189, 211, 213, 252, 280, 281, 346, 454, 455, 589, 6001 646
                  pollutants/pollution: 9, 27, 53, 82, 112, 159, 190, 201, 239, 244, 246, 447, 448, 507, 508, 530, 531, 595, 597,
                         598,618,657








                 A-62
                                                                                                           Bibliography


                 post-larvae: see juveniles
                 predation (by and on summer flounder): 304, 315, 664
                 Raritan Bay: 36, 321
                 recreational fisheries: see fisheries
                 recruitment: 7, 63, 273, 356, 357, 376
                 reproduction: 97, 99, 169, 196, 383, 384, 38@, 397, .199, 424, 446,'561,  651, 656, 579
                 Rhode Island: 204, 225, 264, 265, 320, 456, 496
                 salindy: 141, 328, 329, 334, 335, 336, 470, 471
                 salt marsh habitat: 228, 241, 356, 357, 494, 575, 628, 633, 637
                 salt marsh creek habitat: 64, 76, 77, 121, 494, 516, 518, 519, 522, 553, 612, 614, 630, 631
                 Sandy Hook Bay: 46, 566
                 scallop bed habitat: 542
                 settlement: 63, 284, 383, 492
                 South Atlantic: 16, 17, 34, 175, 425, 510, 599, 615, 634, 635, 636
                 South Carolina: 9, 10, 30, 31, 32, 33, 45, 77, 230, 246, 287, 449, 461, 522, 524, 552, 568, 583, 612, 614, 637,
                       638, 639, 640, 641, 671
                 spawning: see reproduction
                 species list: 26, 176, 177, 183, 197, 214,350, 352, 377, 381, 404, 490, 503, 504, 611, 616, 671
                 survival: see morlality/survival
                 stocks/stock assessment: 81, 130,    134, 135, 144, 175, 207, 221, 269, 270, 406, 497, 607, 617
                 systematics: see identification
                 tagging/marking: 145, 193, 218, 242, 245, 269, 270, 318, 380,393,406
                 temperature: 124, 150, 152, 202, 247, 262, 271, 272, 328, 329, 330, 331, 334, 335, 336, 337, 470,471, 564,
                       593, 594, 602, 605, 621
                 tracking: see movement
                 vegetated habitat: 3, 4, 282, 304, 452, 45.1, 608, 6.10
                 Virginia: 43, 44, 73, 100, 107, 190, 242, 292, 339, 340, 341, 343, 463, 464, 490, 498, 499, 500, 501, 533
                 weight: see biomass, length (incl. length/weight)










               Appendix B
               List of Experts

               Able, Kenneth W., Institute of Marine and Coastal Sciences, Rutgers University, Marine Field Station,
                    800 Great Bay Blvd., Tuckerton, NJ 08087

               Azarovitz, Thomas R., NOAA/National Marine Fisheries Service, Northeast Fisheries Science Center,
                    Woods Hole Laboratory, 116 Water Street, Woods Hole MA 02543-1097

               Bengston, David A., Department of Zoology, University of Rhode Island, Kingston, RI 02 881

               Burke, John S., NOAA/National Marine Fisheries Service, Southeast Fisheries Science Center, Beaufort
                    Laboratory, 101 Pivers Island Rd., Beaufort, NC 28516-9722

               Byrne, Donald M., New Jersey Department of Environmental Protection and Energy, Division of Fish,
                    Game and Wildlife, Nacote Creek Research Station, P.O. Box 418, Port Republic, NJ 08241

               Casey, James F., Maryland Department of Natural Resources, Tidewater Administration, Fisheries
                    Division, 580 Taylor Avenue, Tawes State Off ice Bldg., Annapolis, MD 21401

               Castonguay, Wayne, Massachusetts Division of Marine Fisheries, Cat Cove Lab, Sale m, MA 01970

               Daiber, Franklin Co., College of Madne Studies, University of'Delaware, Newark, DE 19716

               Daniel, Louis B., III Virginia Institute of Marine Science, College of William and Mary, P.O. Box 786,
                    Gloucester Point, VA 23062

               Desfosse, Joseph C., Virginia Institute of Marine Science, College of William and Mary, P.O Box 1046,
                    Gloucester Point, VA 23062

               DeVries, Doug, NOAA/National Marine Fisheries Service, 3500 Delwood Beach Road, Panama City, FL
                    32408

               Epifanio, Charles E., University of Delaware, Marine Biology-Biochemistry Program, 700 Pilottown Rd.,
                    Lewes, DE 19958

               Fahay, Michael P., NOAA/National Marine Fisheries Service, Northeast Fisheries Science Center,
                    James J. Howard Marine Sciences Laboratory, Highlands, NJ 07732

               Fogarty, Michael NOAA/National Marine Fisheries Service, Northeast Fisheries Science Center, Woods
                    Hole. Laboratory, 116 Water Street, Woods Hole MA 02543-1097

               Fonseca, Mark NOAA/National Marine Fisheries Service, Southeast Fisheries Science Center, Beaufort
                    Laboratory, 101 Pivers Island Rd., Beaufort, NC 28516-9721










                 B-2
                                                                                                             Experts

                 Gorski, Stan, NOAAiNational Marine Fisheries Service, Northeast Fisheries Science Center, James J.
                     Howard Marine Sciences Laboratory, Highlands NJ 07731

                 Hettler, William, NOAA/National Marine Fisheries Service, Southeast Fisheries Science Center, Beaufort
                     Laboratory, 101 Pivers Island Rd., Beaufort, NC 28516-9722

                 Hocuff, Charles H., Horn Point Environmental Laboratory, University of Maryland, P.O. Box 775,
                     Cambridge, MD 21613-0775

                 Hoff, Thomas B., National Marine Fisheries Service, Woods Hole Laboratory, 116 Water Street, Woods
                     Hole, MA 02543

                 Howe, Arnold B., Massachusetts Division of Marine Fisheries, Sandwich, MA 02563-1802

                 Howell-Heller, Penelope T., Connecticut Department of Environmental Protection, Bureau of Fisheries,
                    .. P.O. Box 248, Waterford, CT 06385

                 Jearld, Ambrose Jr. NOAA/National Marine Fisheries Service, Northeast Fisheries Science Center,
                     Woods Hole Laboratory, 116 Water Street, Woods Hole MA 02543-1097

                 Jesien, Roman, Horn Point Environmental Laboratory, University of Maryland, PO Box 775, Cambridge,
                     MD 21613


                 Kaiser, Susan C., Institute of Marine and Coastal Sciences, Rutgers University, Marine Field Station,
                     800 Great Bay Blvd., Tuckerton, NJ 08087

                 Keefe, MaryLouise, Oregon Department of Fish & Wildlife, 211 Inlow Hall, Eastern Oregon State
                     College, LaGrande, OR 97850

                 Klein-MacPhee, Grace, University of Rhode Island, Graduate School of Oceanography, South Ferry
                     Road, Narragansett, RI 02882-1197

                 Lynch, Timothy R., Rhode Island Department of Environmental Management, Division of Fish and
                     Wildlife, 150 Fowler St., Wickford, RI 02852

                 Malloy, Kirk D., Boston University, Department of Biology, 5 Cummington Street, Boston, MA 02215

                 Mercer, Linda P., North Carolina Department of Environment, Health and Natural Resources, Division of
                     Marine Fisheries, P.O. Box 769, Morehead City, NC 28557-0769

                 Michels, Stewart F., Delaware Division of Fish and Wildlife, Department of Marine Fisheries, 99 Kings
                     Highway, PO Box 1401, Dover, DE 19903

                 Monaghan, James Patrick, Jr., North Carolina Department of Environment, Health and Natural
                     Resources, Division of Marine Fisheries, P.O. Box 769, Morehead City, NC 28557-0769

                 Murawski, Walter S., New Jersey Department of Environmental Protection and Energy, Bureau of
                     Freshwater Fisheries, CN 400, Trenton, NJ 08625

                 Musick, John A., Virginia Institute of Marine Science, College of William and Mary, P.O Box 358,
                     Gloucester Point, VA 23062









                 Experts                                                                                            B-3





                Norcross, Brenda, University of Alaska- Fairbanks, School of Fisheries and Ocean Sciences, Institute of
                     Marine Science, Fairbanks, AK 99775-1080

                Poole, John C., P.O. Box 167, Melvin Village, NH 03850

                Powell, Allyn B., NOAA/National Marine Fisheries Service, Southeast  Fisheries Science Center,
                     Beaufort Laboratory, 101 Pivers Island Rd., Beaufort, NC 28516-9722

                Powell, Christopher, Rhode Island Department of Environmental Management, Division of Fish and
                     Wildlife, Great Swamp, P.O. Box 218, West Kingston, RI 02892

                Ross, Jeffrey L., North Carolina Department of Environment, Health and Natural Resources, Division of
                     Marine Fisheries, Monteo Field Office, P.O. Box 1550, Manteo, NC 27954

                Rountree, Rodney A., NOAA/National Marine Fisheries Service, Northeast Fisheries Science Center,
                     Woods Hole Laboratory, 116 Water Street, Woods Hole, MA 02543-1097

                Scarlett, Paul G,, New Jersey Department of Environmental Protection and Energy, Division of Fish,
                     Game and Wildlife, Nacote Creek Research Station, P.O. Box 418, Port Republic, NJ 08241

                Shepherd, Gary P., NOAA/National Marine Fisheries Service, Northeast Fisheries Science Center,
                     Woods Hole Laboratory, 116 Water Street, Woods Hole MA 02543-1097

                Shipman, Susan, Georgia Department of Natural Resources, Coastal Fisheries Dept., I Conservation
                     Way, Brunswick, GA 31523-8600.

                Smith, Joseph W., NOAA/National Marine Fisheries Service, Southeast Fisheries Science Center, 101
                     Pivers Island Rd., Beaufort, NC 28516

                Smith, Ronal W., Delaware Division of Fish and Wildlife, 89 Kings Hwy., P.O. Box 1401, Dover, DE
                     19903


                Smith, Walter G., NOAA/National Marine Fisheries Service, Northeast Fisheries Science Center,
                     James J. Howard Marine Sciences Laboratory, Highlands NJ 07732

                Studholme, Anne L., NOAAJNational Marine Fisheries Service, Northeast Fisheries Science Center,
                     James J. Howard Marine Sciences Laboratory, Highlands NJ 07732

                Targett, Timothy E., College of Marine Studies, University of Delaware, 700 Pilottown Rd., Lewes, DE
                     19958


                Terceiro, Mark, NOAA/National Marine Fisheries Service, Northeast Fisheries Science Center, Woods
                     Hole Laboratory, 116 Water Street, Woods Hole, MA 02543-1097

                Wenner, Charles A. South Carolina Wildlife and Marine Resources, P.O. Box 12559, Charleston, SC
                     29412-2559


                Wilk, Stuart J., NOAA/National Marine Fisheries Service, Northeast Fisheries Science Center, James J.
                     Howard Marine Sciences Laboratory, Highlands NJ 07732










                Appendix C
                User Groups

                American Littoral Society
                    Sandy Hook, Highlands, NJ 07732
                    Attention: P. Carlsen


                Connecticut Bureau of Natural Resources, Marine Fisheries
                    P.O. Box 248, Waterford, CT 06385
                    Attention: P.T. Howell, M. Johnson


                Delaware Division of Fish and Wildlife
                    89 Kings Highway, PO Box 1401, Dover, DE 19903
                    Attention: S. Michels, R. Seagrave,

                Harbor Branch Oceanographic Institute, Inc.
                    5600 Old Dixie Highway
                    Ft. Pierce, FL 33946
                    Attention: R.G. Gilmore, Jr.


                Georgia Department of Natural Resources
                    Coastal Fisheries Dept., I Conservation
                    Way, Brunswick, GA 31523-8600.
                    Attention: J.L. Music, Jr., J. Pafford, S. Shipman

                Maryland Department of Natural Resources
                    Tidewater Administration
                    Fisheries Division, 580 Taylor Avenue
                    Tawes State Office Bldg.
                    Annapolis, MD 21401
                    Attention: J.F. Casey

                Massachusetts Division of Marine Fisheries
                    Cat Cove Laboratory, Salem, MA 01970
                    Attention: W. Castonguay

                NOAA/National Marine Fisheries Service
                    Southeast Fisheries Science Center
                    Beaufort Laboratory
                    101 Pivers Island Road
                    Beaufort, NC 28516
                    Attention: J. Burke, S.E. Chester, L. Coston-Clements, D. Hoss, D. Peters, A.B. Powell










                 C-2
                                                                                                             User groups



                 NOAA/National Marine Fisheries Service
                      Northeast Fisheries Science Center
                      James J. Howard Marine Sciences Laboratory
                      Highlands, NJ 07732
                      Attention: A. Bejda, M.P. Fahay, W.G. Smith, A.L. Studholme, S.J. Wilk

                 NOAA/National Marine Fisheries Service
                      Northeast Fisheries Science Center
                      Milford Laboratory
                      Milford, CT 06460
                      Attention: A. Calabrese


                 NOAXNational Marine Fisheries Service
                      Northeast Fisheries Science Center
                      Woods Hole Laboratory
                      166 Water Street, Woods Hole, MA    02543-1097
                      Attention: T. Azarovitz, M. Fogarty, T. Hoff, A. Jearld, B. O'Gorman, G.P. Shepherd, M. Terceiro

                 New Jersey Department of Environmental Protection and Energy
                      Division of Fish, Game and Wildlife
                      Nacote Creek Research Station
                      P.O. Box 418
                      Port Republic, NJ 08241
                      Attention: D. Byrne, P.G. Scarlett, B. Halgren

                 New Jersey Department of Environmental Protection and Energy
                      Division of Fish, Game and Wildlife
                      Bureau of Freshwater Fisheries
                      501 E. State St.,CN 400
                      Trenton, NJ 08625
                      Attention: B. Freeman, W.S. Murawski

                 New York Department of Environmental Conservation
                      Stony Brook, NY 11790
                      Attention: K. McKown, V.J. Vecchio-, B.H. Young

                 State University of New York
                      Marine Science Research Center
                      Stony Brook, NY 11794-5000
                      Attention: J. Schubel


                 North Carolina Division of Marine Fisheries
                      P.O. Box 769, Morehead City, NC 28557
                      Attention: D. DeVries, L. Mercer, J.P.. Monaghan, Jr.

                 North Carolina Division of Marine Fisheries
                      Elizabeth City District Office, Elizabeth City, NC 27909
                      Attention: L. Henry








              User groups
                                                                                                               C- 3




               North Carolina Division of Marine Fisheries
                   Washington District Office, Washington, NC 27889
                   Attention: 0. Moye

               North Carolina Division of Marine Fisheries
                   Wilmington District Office, Wilmington, NC 28402
                   Attention: F. Rohde


               North Carolina Division of Marine Fisheries
                    Monteo Field Office, P.O. Box 1550, Manteo, NC 27954
                   Attention: J.L. Ross


               Rhode Island Division of Fish and Wildlife
                   150 Fowler St., Wickford, RI 02852
                   Attention: T.R. Lynch

               Rutgers, The State University of New Jersey
                   Institute of Marine and Coastal Sciences
                   Marine Field Station, 800 Great Bay Blvd.
                   Tuckerton, NJ 08087
                   Attention: K.W. Able, S.C. Kaiser, D.A. Witting

               South Carolina Wildlife and Marine Resources Center
                   P.O. Box 12559, Charleston, SC 29422-2559
                   Attention: R. Martore, J. Smith, B. Stender, Ft. VanDolah, C.A. Wenner, D.M. Wyanski

               South Carolina Wildlife and Marine Resources Department
                   Waddell Mariculture Center, Bluffton, SC 29910
                   Attention: D. Hamilton, J. Hollaway

               United States Environmental Protection Agency
                   South Ferry Road, Narragansett, RI 02882
                   Attention: D.A. Bengsten

               United States National Park Service
                   Biscayne Bay National Park, Homestead, FL 33030
                   Attention: T. Rutledge

               University of Delaware
                   College of Marine Studies
                   700 Pilottown Road
                   Lewes, DE 19958
                   Attention: C.E. Epifanio, T.E. Targett

               University of Maryland
                   Horn Point Environmental Laboratory
                   University of Maryland System, Cambridge, MD 21613
                   Attention: C. Hocutt, R. Jesien










                  C-4
                                                                                                          User groups



                  University of Massachusetts
                      Cooperative Extension, Essex County Office
                      562 Maple Street
                      Hawthorne, MA 01937-0362
                      Attention: S. Jacobson

                  University of North Carolina
                      Wilmington, NC 28402
                      Attention: D. Lindquist

                  University of North Florida
                      'Coastal Fisheries Laboratory, Jacksonville, FL 32203
                      Attention: C. DeMort


                  University of South Carolina
                      Belle Baruch Marine Laboratory, Georgetown, SC 29440
                      Attention: D. Allen, G. Ogburn

                  University of South Carolina
                      Coastal Carolina College
                      P.O. Box 1954, Conway, SC 29526
                      Attention, R. Moore


                  Virginia Institute of Marine Science
                      College of William and Mary
                      P.O. Box 1046, Gloucester Point, VA 23062
                      Attention: J.C. Desfosse, J.A. Musick


































                                          U.S. GOVERNMENT PRIN71NG OFFICE: 1994 - 3 0 0 - 5 6 6 1 0 4 0 4 0



























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                                                                Life History of Summer Flounder. Adults generally spend summer in estuaries and on
                                                                the shallow continental shelf, migrate offshore in autumn, spend winter in the ocean, and
                                                                migrate back inshore in spring. Adults spawn in the ocean during autumn and winter.
                                                                Eggs rise to near-surface waters and larvae hatch. Larvae are pelagic and begin
                                                                development in the ocean but they enter estuaries during winter and spring to complete
                                                                development and settle. Juveniles grow rapidly during the summer and join adults in
                                                                offshore migration in autumn, returning to estuaries the following year.































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