Cover il/uslrafr PI: L@ His tory of Sz~nzmer Hourader 

(/czrger view expfni~led in detail 011 inside hack cover) 

This publication should be cited as: 

Able, Kenneth W. and Susan C. Kaiser. 1994. Synthesis of Summer Flounder Habitat Parameters. NOAA 

Coastal Ocean Program Decision Analysis Series No. 1. NOAA Coastal Ocean Office, Silver Spring, MD. 

68 pp. + biblio. + 3 apps. 

This publication does not constitute an endorsement of any commercial product or 

intend to be an opinion beyond scientific or other results obtained by the National 

Oceanic and Atmospheric Administration (NOAA). No reference shall be made to 

NO& or this publication hshed by NOAA, in any advertising or sales promotion 

which would indicate or imply that NOAA recommends or endorses any proprietary 

product mentioned herein, or which has as its purpose an interest to cause directly or 

indirectly the advertised product to be used or purchased because of this publication. 
eaders 

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 

resources, can make significant strides toward finding solutions to critical problems. By working 

together toward these solutions, we can ensure the sustainability of these coastal resources and 

allow for compatible economic development that will enhance the well-being of the Nation now 

and in future generations. The goals of the program parallel those of the NOAA Strategic Plan 

for 1995-2005. 

A specific objective of COP is to provide the highest quality scientific information to coastal 

managers in time for critical decision making and in a format usehl for these decisions. To help 

achieve this, COP inaugurated a program of developing documents that would synthesize 

information on issues that were of high priority to coastal managers. To develop such documents, 

a three-step process was used: 1) to compile a list of critical topics in the coastal ocean through 

a survey of coastal resource managers and to prioritze and select those suitable for the document 

series through the use of a panel of multidisciplinary technical experts; 2) to solicit proposals to 

do research on these topics and select principal investigators through a rigorous peer-review 

process; and 3) to develop peer-reviewed documents based on the winning proposals. 

Seven topics and associated 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. 

Therefore, we encourage you to write, fax, call, or Internet us with your comments. Please be 

assured that we will appreciate these comments, either positive or negative, and that they will help 

us direct our future efforts. Our address and telephone and fax numbers are on the inside front 

cover. My Internet address is DSCAVIA@HQ.NOAA.GOV. 

/~onald Scavia 

Director 

NOAA Coastal Ocean Program 

Table of Contents 

LIST OF FIGURES AND TABLES 

ACKNOWLEDGMENTS 

EXECUTIVE SUMMARY 

CHAPTER 1:INTRODUCTION 

Background and Overview 

Scope 

CHAPTER 2: DISTRIBUTION AND LIFE HISTORY PATTERNS 

Geographical Distribution 

General Life History 

Stock Identification 

CHAPTER 3: EGGS 

lntroduction 

Spatial and Temporal Distribution 

Temperature 

CHAPTER 4: OFFSHORE LARVAE 

lntroduction 

Spatial and Temporal Distribution 

Temperature 

Substrate 

CHAPTER 5: ESTUARINE LARVAE 

lntroduction 

Spatial and Temporal Distribution 

Temperature and Salinity 

Substrate and Behavior 
iv 

Table of Contents 

CHAPTER 6: ESTUARINE JUVENILES 

lntroduction 

Spatial and Temporal Distribution 

Temperature, Salinity and Dissolved Oxygen 

Substrate 

Behavior 

CHAPTER 7: OFFSHORE JUVENILES 

lntroduction 

Spatial and Temporal Distribution 

Temperature 

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 

APPENDICES 

Appendix A. 

Comprehensive Summer Flounder Bibliography 

Author Index 

Subject l ndex 

Appendix B. 

List of Experts 

Appendix C. 

User Groups 
List of Figures and Tables 

Fisures 

East coast of North America location maps showing summer flounder range. 

Cumulative egg distribution and abundance over northern and mid-Atlantic 

continental shelf. 

Egg abundance by month and region over the north and mid-Atlantic 

continental shelf. 

Monthly larval distribution and abundance over the north and mid-Atlantic 

continental shelf. 

Larval abundance by month and region over the north and mid-Atlantic 

continental shelf. 

Transformation stages of summer flounder. 

Locations of summer flounder studies: (A) Great Bay-Little Egg Harbor, New 

Jersey; (B) Charleston Harbor, South Carolina; (C) Long Island Sound, New 

YorkIConnecticut; and (D) Pamlico-Albemarle sounds, North Carolina. 

Cumulative length- and stage-frequency distributions for transforming larvae in 

two marsh creeks in Great Bay-Little Egg Harbor. 

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

Harbor. 

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

Synopsis of seasonal movements from tagging studies. 

Seasonal variation in abundance of transforming larvae in Little Sheepshead 

Creek, Great Bay-Little Egg Harbor. 
Figures and Tables 

Seasonal variation in abundance of transforming larvae in Schooner Creek, 

Great Bay-Little Egg Harbor. 

Monthly length-frequency distributions for juveniles in four marsh creeks in 

Charleston Harbor. 

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

Sound. 

Monthly distribution and abundance of juveniles and adults in Long Island 

Sound. 

Seasonal length-frequency distributions for juveniles and adults in PamlicoAlbemarle sounds. 

Monthly length-frequency distributions for juveniles in shallow marsh habitats in 

Pamlico-Albemarle sounds. 

Seasonal distribution and abundance of juveniles and adults in PamlicoAlbemarle sounds. 

Bimonthly length-frequency distributions for juveniles in Charleston Harbor 

Juvenile abundance relative to salinity in four marsh creeks in Charleston 

Harbor. 

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

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

shelf. 

Sampling locations for two inner continental shelf trawl surveys in the South 

Atlantic Bight. 

Monthly length-frequency distributions for juveniles and adults on the South 

Atlantic Bight inner continental shelf. 

Monthly distribution of adults on the north and mid-Atlantic continental shelf 

from the commercial fishery. 

July length-frequency distributions of adults in Great Bay from the recreational 

fishery over a 10-year period. 
Figures and Tables 

Tables 

6.1 Characteristics of marsh creek habitat for juveniles in Charleston Harbor 

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

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 Ill (Virginia lnstitute of Marine Sciences) for habitat data on juveniles in 

marsh creeks; J. Musick and J. Desfosse (Virginia lnstitute 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 Division of Fish, Game and Wildlife); P. Carlsen 

(American Littoral Society); J. Casey (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. 

Gilmore (Harbor Branch Oceanographic Institute); T. Hoff (Mid-Atlantic Marine 

Fisheries Council); D. Hoss (National Marine Fisheries Service, Beaufort Laboratory); 
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. 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 Program, and Rutgers University, Institute of Marine and 

Coastal Sciences (IMCS). This is 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 reflected in the compressed age structure of the 

population and the low catches in both commercial and recreational fisheries. 

Declining habitat quantity and quality may be contributing to these declines, however 

we lack a thorough understanding 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 f IOU nder (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 
xii 

Executive Surnrnarv 

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 completed 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 synthesis 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 Surnrnaw 

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 flounder in 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 

juveniles and adults, although the depth of understanding varies from region to region. 

Due to the highly migratory nature of 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 recommend that estuarine juveniles should be the focus of researchers 

and resource managers because: 1) juvenile grow h and 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 marsh creeks and shallow portions of bays 

appear to be most important as nurseries, especially for the early juvenile stages. We 

therefore suggest that resource managers pay particular attention to these habitats 

and their maintenance in order to improve the status of summer flounder populations. 

Chapter 1 

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 

ServiceINortheast 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; Szedlmayer 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 
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 

1971'). 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 effects). We have 

cautiously interpreted prior generalizations regarding summer flounder habitat, and are 

particularly careful not to extrapolate from general statements concerning other closely 

related flounders (Paralichthys spp.). Because pertinent life history information is 

frequently lacking, we include a brief summary of our current understanding of the 

salient aspects. Since different stocks or populations may have different habitat 

requirements or patterns of habitat use, we also briefly summarize what is known 

about distribution and stock identification. We recognize that we still lack important 

information about the habitat of summer flounder during 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 Azarovitt 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 bycatch. 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 
Distribution 

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

mentioned in the text. 
Distribution 

Figure 2.2. Cumulative distribution and mean abundance of summer flounder eggs from National Marine 

Fisheries Service (NMFS), Marine Resources Monitoring, Assessment and Prediction (MARMAP) 

offshore surveys during 1977-1 964 (Sherman 1980, 1986). tdonthly to bimonthly plankton samples were 

collected from Cape Sable to Cape Hatteras (Fig. 2.1) using 61 cm bongo frames (Sibunka and 

Silverman 1984; Morse et al. 1987). The 200 m and 1000 m contours are shown. (Figure from Able et 

al. 1990). 
Distribution 

100 GEORGES BANK 

DELMARVA PENNINSULA 

NS 

VIRGINIA CAPES 

TO CAPE HATTERAS 

NS 

I I 1 I I I I I I I I 

JUL AUG SEP OCT NOV DEC JAN FEB 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 

44 76 74 72 

Paralichthys dentatus 

1977-1987 

Larvae / 10 m2 

44 75 74 72 

Paralichthys dentatus 

1977-1987 ... '. Larvae / 10 mz J.. './ . .'. 

7,,,, 

44 76 14 $2 ~~9.6~ ' iia 44 

Paral~chthys dentatus w*, dfl 

Larvae 1 10 mz ,, .' 

Figure 2.4. Monthly distribution and mean abundance of summer flounder larvae from National Marine 

Fisheries Service (NMFS), Marine Resources Monitoring, Assessment and Prediction (MARMAP) 

offshore surveys during 1977-1 987. Monthly to bimonthly plankton samples were collected from Cape 

Sable to Cape Hatteras (Fig. 2.1) using 61 cm bongo frames (Sibunka and Silverman 1984; Morse et al. 

1987). The 200 m contour is shown. (Figure by M.P. Fahay, National Marine Fisheries Service, Sandy 

Hook, New Jersey). 
Distribution 

14 76 74 72 

Paralichthys dentatus 

1977-1987 

44 74 72 

Paralichthys dentatus 

1977-1987 

Figure 2.4 (continued). 
Distribution 

100 3 GEORGES BANK 

SOUTHERNNEWENGLAND 

NEW JERSEY 

100 

100 DELMARVA PENNINSULA 

VIRGINIA CAPES 

TO CAPE HATTERAS 

NS - 

I I 1 1 I I I I I 1 I I 

JUL AUG SEP OCT NOV DEC JAN FEB MAR APR MAY 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. 
10 

Distribution 

between Chesapeake Bay and Georges Bank. During November and December, 

when the larvae are most abundant in every region, they are fairly evenly distributed 

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 remaining larvae are concentrated off North Carolina. 

As the larvae undergo eye migration during transformation (Fig. 2.6), they enter 

estuarine nursery areas (Burke et al. 1991; Keefe and Able 1993). Based on data from 

the Great Bay-Little Egg Harbor estuarine 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 through 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 late larva 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 eye migration, 

pectoral fin and scale formation be referred to as transforming larvae, as Burke (1991) 

has done. If larvae enter the estuary when water temperatures are low, mortality can 

occur (Szedlmayer 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 et al. 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 (Szedlmayer 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; Szedlmayer 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 estuaries and onto the continental shelf (Able et al. 

1990; Szedlmayer and Able 1993). The adults follow a similar pattern (Fig. 2.10, 2.1 l), 

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. 
Pre-transformation F - 

Figure 2.6. Classification of transformation stages for summer flounder based on degree of e

(adapted from Keefe and Able 1993). The right and left eyes are bilateral and symmetrical in pre-tr

individuals. At the first stage of transformation, F -, the eyes are bilateral but asymmetrical with the 

dorsal to the left eye. By Stage G, the right eye is visible from the left side of the fish. Stage H - diff

that the cornea of the eye is visible from the left side of the fish. At Stage H, the right eye has reach

midline. By Stage H +, the right eye has reached the left side of the head but has not yet reached its 

place. At Stage I , the eye is set in the socket and the dorsal canal has closed. 
I Harbor I 

Distribution 

Figure 2.7. A. Great Bay - Little Egg Harbor 

estuarine system in southern New Jersey with 

the 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 

Sound system bordered by New York and 

Connecticut: D. Pamlico-Albemarle sounds 

estuarine system in North Carolina. 
Distribution 

SHEEPSHEAD 

NOV 1990 - NOV 1991 

80~ SCHOONERCREEK 

Feb 1990 - Apr 1992 

n = 77 

.. 8-8.9 10-1 0.9 12-1 2.9 14-1 4.9 

STANDARD LENGTH (mml 

SCHOONER CREEK 

Feb 1990 - Apr 1992 

n =51 

60 

TRANSFORMATION STAGE 

Figure 2.8. A. Length-frequency, and, B. stage-frequency distributions for 

transforming summer flounder larvae collected during Rutgers University 

Marine Field Station ichthyoplankton surveys in two Great 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 1 mm mesh. See Fig. 2.6 for 

transformation stage descriptions. 
Distribution 

30: - 

NOV - APR 21 - 26 JUL 

20' n = 65 n =21 - 

10: - 

O-lllirll,l ll,lll, 

30: 18 - 27 MAY 2- 10AUG 

20; n=7 n = 106 

101 

0- '7 I 1, I I I I I,, I I,, 

14- 15 JUN 17 - 25 AUG 

n = 114 n = 125 

II~III~SII~ 

28 - 30 JUN 30 AUG - 08 SEP 

n = 38 n = 92 

17- 22 SEP 

n = 50 

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 E g 

Harbor estuarine system (Fig. 2.7A; adapted from Szedlmayer et al. l9&7. 
Distribution 

11 to 25 

SPRING 

Fig. 2.10. Seasonal distribution and abundance of summer flounder from 

National Marine Fisheries Service (NMFS), Northeast Fisheries Science 

Center (NEFSC) groundfish surveys for spring (1982-1990) and autumn 

(1 981 -1 990). 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). 
Distribution 

Figure 2.1 1. Synopsis of results of tagging studies on adult summer 

flounder depictmg, A. offshore movement from summer feeding grounds 

and, B. inshore movement from overwintering grounds. 0 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 followin sources: 

Westman and Neville 1946); Hamer and Lux (1 962); Poo e (1 962); r 9 

Murawski (1 97Oa); Scar ett (1 984); Mercer et al. (1 987); Desfosse et al. 

(1 990); Jesien and Hocutt (1 991). 
Distribution 

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 interpretation 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 transHatteras 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 probably not 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 

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 15°C 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 - 22.9OC, with most occurring in water temperatures 

of 13 - 17.9OC (Smith 1973a,b). Laboratory studies have shown that egg incubation 

time and water temperature are positively correlated between 5 - 21°C, 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 oncethrough cooling systems of power plants (Smith et al. 1979). Late embryo stage eggs 

were exposed to sudden increases in temperature (pre-experiment acclimation = 

16°C; experimental shock exposures = 23, 26, 29, 32, and 35°C) for time periods 

ranging from 3 - 180 minutes at each shock level, and survivorship was high. One 

hundred percent mortality occurred only at 35OC 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 the presence of the larvae of two and sometimes three 

other species of Paralichthvs (Powles and Stender 1976). 

TEMPERATURE 

Larvae have been collected over a wide range of water temperatures (0 - 23. 1°C) 

between Cape Cod and Cape Lookout, with most occurring at temperatures of 9.0 - 

17.9OC (Smith 1973a,b). In the laboratory, yolk sac larvae died at temperatures below 

1 1°C, but those held above 11°C 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 +lO°C thermal shock 

(base 15.1°C; Deacutis 1978), however mortality resulted from shock exposure to 32°C 

(Hoss et al. 1974). 
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; Szedlmayer 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 l nlet 

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; 
Estuarine Larvae 

Y' 

E 

I? 

a 

3 

0 

s 

z 

LL 

0 

a 

w 

m 

z 

3 

z 

Z 

i.5 

5 

Figure 5.1 

0.012 

0.004 

0.000 , 

OCT NOV DEC JAN FEB MAR APR MAY JUN 

. 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 t~des. 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 

1'2 34'5'1 2'3'4'5'1'2 34 5'1 2'3 4'5'1 '2'3'45'1 28 4'5'1 '2.3' 

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 ichthyoplankton 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. 
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 Paralichthvs 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 - 

13°C in Middle Atlantic Bight estuaries (Olney and Boehlert 1988; Wyanski 1990; 

Szedlmayer et al. 1992), from 8.4 - 23.4"C in South Carolina (McGovern and Wenner 

1990) and from 2 - 22°C in North Carolina (Williams and Deubler 1968a). Larvae 

collected in Little Sheepshead Creek (Figs. 2.7, 5.1) typically occurred at temperatures 

of 0 - 12.0°C. In Schooner Creek (Figs. 2.7, 5.2), larvae were collected at -2.0 - 14°C. 

In the laboratory, however, water temperatures less than 2°C killed transforming larvae 

(Szedlmayer et al. 1992). In another laboratory study of the same population, mortality 

of mid-transformation larvae held at 4°C was significantly greater than those held at 

10°C (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 17°C to 93 days at average temperatures of 6.6"C. 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 Paralichthvs responded to die1 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 

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 1968a,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 

 st-uarine 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; 

Szedlmayer 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 TL) 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 TL) 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; Szedlmayer 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 
Estuarine Juveniles 

NOVEMBER 80 APRIL 

n=O n = 143 

@j Lachicotte 

eresford 40 

0 Inlet 

0 

20 DECEMBER 20 MAY n=O n=31 

10 10 

Y 

a .O 0 .. 

3 

n 

20 JANUARY 20 JUNE 

- n=25 

n n=87 

z 

LL 10 10 

0 

u 

m Wo 0 

E 5 160 FEBRUARY 20 

n = 245 JULY 

n = 1 

80 10: 

0 o;... m.., 

80 MARCH 20 

n = 134 

40 10 

0 

AUGUST 

n=O 

TOTAL LENGTH (mm) 

Figure 6.1. Length-frequency distributions for juvenile summer flounder collected 

during 1986 (November, December) -1 987 from four Charleston Harbor estuarine 

system marsh creeks (Fig. 2.7B) using a rotenonelblock net method (Wenner et 

al. 1990). 
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. 2.7~) 

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 falliearly winter 

(November), they have moved offshore into the Atlantic Ocean. In Great Bay, youngof-the-year spend the summer in the estuary and begin leaving as early as August 

(Able et al, 1990; Rountree and Able 1992a; Szedlmayer and Able 1992; Szedlmayer 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). 
Estuarine Juveniles 

161 1 APRllL 1 APRIL 

MAY 

n = 41 

JUNE 

n = 7 

SEPTEMBER 

12 n = 70 

SEPTEMBER 

n=49 

OCTOBER OCTOBER n = 101 n = 55 

20 

NOVEMBER k NOVEMBER 

12 n = 28 n = 1 

TOTAL LENGTH (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 

rn otter trawl at, typicall , 40 stations that were chosen by stratified random 

design. Data are baseJon the finfish surveys of the Connecticut Division of 

Marine Fisheries (1 99Oa, 1990 and 1992). 
Estuarine Juveniles 

Figure 6.3. Distribution 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 

1 992). 
Estuarine Juveniles 

Figure 6.3 (continued). 
" = 710 MARCH 

J 

JUNE 

n = 920 n JUNE = 152 

1 

U SEPTEMBER 

SEPTEMB

n = 535 

I 

3 40 

0 

DECEMBER 

n= 115 

DECEMBE

NO SAMP

TOTAL LENGTH (mm) 

Figure 6.4. Length-frequency distributions for juvenile and adult summer flounder in the Paml

sounds estuarine system (Fig. 2.7D) during years of high (1 987) and low (1 990) abundance. A stra

design was used to choose sites throughout Pamlico Sound, the lower reaches of Albemarle Sou

subestuaries (Pamlico, Neuse and Pungo rivers) for sampling by 20 minute tows of a Mongoose tra

months in 1987 and 3 months in 1990. Data are based on the North Carolina Division of Ma

(1 987-1 991) trawl survey. 
Estuarine Juveniles 

1 I987 (n = 185) 

16 

12 JUN 

8 I987 n =75 

1990 n = 24 

4 

0 

Y 12 JUL 

8 1987 n =27 3 1990 n = 21 

0 - 4 

U. 

0 1987 n = 20 

CT 1990n=13 w4 

m 

E 0 

1987 n = 19 

1990 n = 1 

4 = 0 

161 

OCT 

1987 n = 13 

1990 n =6 

4 

NOV 

1987 n=3 

4 1990 NS 

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. 2.7D) 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 rn seines. Data 

from Noble and Monroe (1 991). 
Estuarine Juveniles 

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 (1 987-1 991) trawl survey. 
Estuarine Juveniles 

Figure 6.6 (continued). 
Estuarine Juveniles 

JANUARY 

n = 67 

MARCH 

n = 136 

30 - 

V) MAY 

J 

a 20- n = 161 

z 

2 

a 30- 

w JULY 

rn 3 20- n = 162 

z 

10- 

0-r'. r.g-. 

SEPTEMBER 

n = 23 

NOVEMBER 

n = 74 

TOTAL LENGTH (cm) 

Figure 6.7. Length-frequency distributions for juvenile summer flounder 

in channels in the Charleston Harbor estuarine system (Fig. 2.78) 

during 1985-1988. Bimonthly sampling was conducted using 4.9 and 

7.6 semiballoon 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.) 
Estuarine Juveniles 

TEMPERATURE, SALINITY AND DISSOLVED OXYGEN 

Estuarine juveniles can be found in a wide range of naturally occurring 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 - 14°C) 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 
Table 6.1. Physical characteristics of four marsh creek study sites in the Charleston Harbor estuary (Fig. 6.1) 

for 13 months from June 1986 - August 1987 (Hoffman 1991). Fishes were collected by setting paired blo

and downstream of the sites, releasing rotenone upstream, and then dip-netting and seining. See Fig

locations; 1 = Pita Creek; 2 = Lachicotte Creek; 3 = Beresford Creek ; 4 = Inlet Creek. (mlw = mean low wate

(SITE NO.) 

LOCATION 

DISSOLVED TEMPER- SALINITY OXYGEN DEPTH 

ATURE SUBSTRATE 

(OC) (PP~) (PP~) (m) 

(1) 35.6 km from 7.6-32 0.8-24.5 4.2-10.2 0.6 - 1.1 at mlw shell hash 

mouth of Charleston and sand 

Harbor 

(2) 30.7 km from 7.3-30.9 7.1 -25.2 1.6-1 1.2 0.7 - 1.8 at mlw sand and shel

mouth of Charleston hash with som

Harbor mud downstre

(3) 20.7 km from 9.6-31.5 13.0-23.7 2.0-1 1.7 0.4 at mlw mud upstream

mouth of Charleston sand and som

Harbor shell downstre

(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 

mouth of Charleston very little mud

Harbor in a creek off small oyster b

the intracoastal in upper 112 p

waterway ially exposed a

low tide 
Estuarine Juveniles 

JANUARY 

n=18 

FEBRUARY 

60 n = 245 

40 

20 

APRIL 

n = 143 

MAY 

n=31 

SALINITY (ppt) 

20 

40/.. , , , n, a 

. , . ,, . r20 

0- 

Figure 6.8. Abundance of juvenile summer flounder 

relative to salinity in four Charleston Harbor estuarine 

system marsh creeks (Fig. 2.7B) collected by a 

rotenone 1 block net method during 1987 by the South 

Carolina Marine Resources Center. Data based on 

Wenner et al. (1 990). 

80 - 

60. 

40. 

20. 

JULY 

n-1 
(SITE NO.) 

LOCATION 

Table 6.2. Physical characteristics of four marsh creek study sites in lower Chesapeake Bay and the ocean sid

shore of Virginia (Fig. 2.1) sampled monthly from June - November in 1989 and May - November in 1990 (Dan

Fishes were collected by setting paired block nets upstream and downstream of the sites, releasing rotenone u

then dip-netting and seining. 

MEAN 

TEMPER- SALNITY 

ATURE LOW WATER SUBSTRATE VEGETATION 

(PPV (OC) (m) 

(1) small tributary 

of the Poropotank River; 

-3.1 -6.1 m wide, -30.5 m 

long 

(2) in Goodwin Islands 

at mouth of York River; 

-9.1 m wide, -30.5 m long 

(3) creek off main channel 

entering town of Wachapreague; -6.1 m wide, 

-30.5 m long 

(4) behind Parramore 

Island, close to mouth 

of Little Machipongo 

Inlet; -6.1 m wide, -30.5 

m long 

15-31 3-1 1 0.3-0.5 shell hash and off Spartina 

very muddy marsh, some 

Juncus 

16-30 1 5-24 0.9 hard-packed mud off SDartina 

and shell hash in marsh; Sparkin

center, very island in 

muddy sides center 

15-30 26-29 0.8-0.9 shell hash in cen- none 

(hole= ter and steep 

1.1 -1.2) muddy sides 

17-28 28-33 0.15-0.2 sandy mud off Spartina 

(hole=1.2) marsh 
Estuarine Juveniles 

SPRING 

10 (MAY - JUNE) 

- I TRIAL I TRlAL ll I TRlAL l TRlAL ll I TRIAL 1 TRIAL II I 

AUTUMN 

(NOVEMBER - 

DECEMBER) 

EELGRASS 

SEA LETTUCE 

UNVEGETATED 

0 MUD 

I TRlAL l TRIAL ll 1 TRlAL l TRlAL ll I TRlAL l TRIAL 11 1 

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 (1 992). 
Estuarine Juveniles 

45 

stage as had been observed in laboratory experiments (Keefe and Able 1994). At 

temperatures below 2 - 3OC, significant mortality has been reported in laboratory 

experiments (Malloy 1990; Malloy and Targett 1991; Szedlmayer 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-ofthe-year juveniles (Szedlmayer 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 - 

28,0°C), 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 - 24.9"C), 

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 f 1.2 mm SL) preferred sand substrate (< 5% 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 (01th 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 
Estuarine Juveniles 

settled and small juveniles (17 - 41 mm SL) based on caging experiments was variable in 

spring (range 0.18 - 0.89 mmlday). 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 die1 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 

die1 cycle of movements in the 1-km-long Schooner Creek (Fig 2.7a; Szedlmayer and 

Able 1993). These young-of-the-year individuals (210 - 254 mm TL) 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 

mantlma) 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 autumn 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 well-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 (I00 - 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 
Offshore Juveniles 

AUTUMN 1991 

age = 0 

n = 178 

** : '/ . . *( -. 

. 

.. 

... . WINTER 1992 

;$-,,:, 8; age = 1 

.C !.., / n = 627 .' ,. 

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 (N 2 FSC) 

groundfish surveys for A, autumn, B. winter, and C. s ring, 1991 - 1992. Sampling was 

conducted by stratified random design in the Gulf of haine 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 1000 m contour is shown. Collections 

where no 'uveniles were caught are omitted. (Adapted from ma s provided by G. 

Shepherd. tbationai Marine Fisheries Service. Woods Hole Laboratory. hassachusetts). 
Offshore Juveniles 

mlico Sound 

arleston Harbor 

Helena Sound 

ockwood Folly Inlet 

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 6. 

sample areas for two South Atlantic Bight inner continental shelf (c 10 m) trawl 

surveys. Wenner and Sedberry (1989) sampled at randomly chosen sites within each 

stratum during 1980, 1981 and 1982. Beatty et al. (1 989) sampled four 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. 
Offshore Juveniles 

JANUARY JULY 

30 n = 177 

(NO SAMPLING 

20 . FOR STUDY B) 

10. 

o..,.... 

50 - 

40 - FEBRUARY 

30 - (NO SAMPLING 

FOR EITHER 

20 - STUDY A OR 8) 

10 - 

0 0 . . . . . . . . , , , . . . ., . ., , , , . , . . . . . . . . . , . , . . 

MARCH SEPTEMBER 

n = 282 

(NO SAMPLING 

FOR EITHER 

20 

STUDY A OR B) 

2 10 n 

z - 0 

& 50: 

40- APRIL 

n = 185 OCTOBER 

(NO SAMPLING 

FOR STUDY B) 

MAY / 

n = 253 

NOVEMBER 

n = 305 

JUNE - 

n=106 : 

DECEMBER 

n = 22 

(NO SAMPLING FOR 

STUDY A: FEW 

STUDY B SAMPLES) 

10 

0 I"'."-+ 

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 Sedberty 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 

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 15°C (Sissenwine et al. 1979), but 

nothing else is known primarily because juveniles and adults are not differentiated. 

Chapter 8 

~stuarine 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 
JANUARY I 

APRIL 

FEBRUARY 

MAY 

MA

Figure 8.1. Average annual landings (metric tons per ten minutes square of latitude and longitude) of summer flounder captured in th

States commercial fishery during 1987-1989 from the Gulf of Maine to off Cape Hatteras (Fig. 2.1) by month. The 200 m contou

(Adapted from maps provided by G. Shepherd, National Marine Fisheries Service, Wocds Hole Laboratory, Massachusetts.) 
Adults 
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 lsland, 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 m 

(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 lsland 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 lsland Sound, individuals that are presumably age I+ 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 lsland 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 

TOTAL LENGTH (cm) 

Figure 8.2. Length-frequency distributions of summer flounder in Great Bay (Fig. 

2.7A) 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). 
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 Paralichthvs 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 (c 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 - 19°C (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-ofthe-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; 1991a,b; Burke 199 1; Hoffman 1991; Noble and 

Monroe 1991; Szedlmayer et al. 1992). 

Summer flounder oxygen requirements have not been well studied, however some 

reports indicate that episodes of hypoxia (low dissolved oxygen, -c 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 beststudied 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 

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; Smith 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 

Schroeder 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 - 3°C) 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 hypoxiaianoxia 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 
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 estuarinedependent 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 influence 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 ("C): Temperature in degrees Celsius. To convert to Fahrenheit, use equation OF = (915 x "C) + 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 shell, 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 die1 patterns. 

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

respiration. 

Ebb tide: Tidal stage at which ocean water flows in an offshore direction and estuarine waters flow 

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

unfertilized state. 

Electrophoresis: 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 lefteye 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 life 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 in some 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. 
Glossary 

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 andlor 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 andlor 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. 
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. 

Paralichthys 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). 
Glossary 

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). 

Scophthalmus 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 1.0 - 2.0 mm diameter (very coarse) to 0.062-0.125 mm 

in diameter (very fine) based on the Wentworth scale. 

Silt: 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 

intertidal 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 marine 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. 
Glossary 

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-0 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 

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. 

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(1): 1-12. 

Adams, S.M. 1976a. The ecology of eelgrass, Zostera marina (L.), fish communities. I. Structural 

analysis. Journal of Experimental Marine Biology and Ecology 22: 269-29 1. 

-- . 1976b. The ecology of eelgrass, Zostera marina (L.), fish communities. ll. Functional 

analysis. Journal of Experimental Marine Biology and Ecology 22: 293-31 1. 

Agnello, R.J. 1989. The economic value of fishing success: an application of socioeconomic 

survey data. Fishery Bulletin, U.S. 87(1): 223-232. 

Ahlstrom, E.H., K. Amoaka, D.A. Hensley, H.G. Moser and B.Y. 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. 

Allen, D.M. and D.L. Barker. 1990. Interannual variations in larval fish recruitment to estuarine 

epibenthic habitats. Marine Ecology Progress Series 65: 1 13-125. 

Allen, D.M., J.P. Clymer Ill and S.S. Herman. 1978. Fishes of the Hereford Inlet estuary, 

southern New Jersey. Lehigh University and The Wetlands Institute. 138 pp. 

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. 

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 lnstitute Special Publication No. 82-1. 

University of South Carolina, Columbia. 275 pp. 

Almeida, F.P., RE. Castaneda, R.V. Jesien, R.C. Greenfield and J.M. Burnett. 1992. Proceedings 

of the NEFCIASMFC summer flounder, Paralichth~s dentatus, aging workshop; 11-13 June 1990, 

Northeast Fisheries Center, Woods Hole, Massachusetts. NOAA Technical Memorandum 

NMFS-FINEC-89. National Marine Fisheries Service. 
Bibliography 

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16-17. 

Ames, W.H. 1954. Biological survey of the Delaware River estuary. Biennial Report 1953-1954, 

2. University of Delaware, Marine Laboratory. pp. 21-31. 

Anderson, E.D., J.M. Mason, 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. 

Anderson, V.T., Jr. 1978. Reverse summer flounder (Paralichthys dentatus) from the Middle 

Atlantic Bight. Bulletin of the New Jersey Academy of Science 23(1): 39-41. 

Anderson, W.W. 1968. Fishes taken during shrimp trawling along the South Atlantic coast of the 

United States, 1931-35. Special Scientific Report-Fisheries 570. United States Fish and Wildlife 

Service. 60 pp. 

Anderson, W.W. and J.W. Gehringer. 1965. Biological-statistical 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. 

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

summer of 1949. Copeia 1951. 

Arve, J. 1960. Preliminary report on attracting fish by oyster-shell plantings in Chincoteague 

Bay, Maryland. Chesapeake Science 1: 58-65. 

Atlantic States Marine Fisheries Commission Subcommittee for Fisheries of the Atlantic Bight. 

1966. A summary of current research on the summer flounder. 2 pp. 

Atlantic States Marine Fisheries Commission Summer Flounder Subcommittee. 1970. Life history 

of the summer flounder. A report to the Advisory Committee. 10 pp. 

. 1971. Life history of the summer flounder. A report to the Advisory Committee. 8 pp. 

Azarovitz, T.R., C.J. Byrne, E.S. Bevacqua, L.I. Despres and H.A. Foster. 1980. Distribution and 

abundance trends of 22 selected species in the Middle Atlantic Bight from bottom trawl surveys 

during 1967-1979. Final report to BLM, AA550-lA7-35. National Marine Fisheries Service. 

Azarovitz, T.R., C.J. Byrne, E.S. Pritchard, L.I. Despres-Patanjo and H.A. Foster. 1985. 

Distribution and abundance trends of 22 selected species in the Middle Atlantic Bight from bottom 

trawl surveys during 1967-1979. Final Report to United States Mineral Management Service, 

Contract No. AA 55GlA7-33. National Marine Fisheries Service, Woods Hole Laboratory, 

Massachusetts. 

Azarovitz, T.R. and M.D. Grosslein. 1987. Chapter 30: Fishes and squids. In Backus, R.H. and 

D.W. Bourne, Editors. Georges Bank. Massachusetts Institute of Technology, Cambridge, 

Massachusetts. 
Biblioaraphy 

Bailey, R.M., J.E. Fitch, E.A. Herald, E.A. Lachner, C.C. Lindsey, C.R. Robins and W.B. Scott. 

1970. A list of common and scientific names of fishes from the United States and Canada. 

Special Publication 6. American Fisheries Society. 150 pp. 

Baptist, J.P. and T.J. Price. 1962. Accumulation and retention of cesium-137 by marine fishes. 

Fishery Bulletin, U.S. 62: 177-187. 

Bass, D.G., Jr. 1983. North Florida streams research project, Study Ill: Rivers of Florida and their 

fishes. Completion Report for Investigations Project, Dingell-Johnson F-36. Florida Game and 

Fresh Water Fish Commission, Tallahassee. 

Bean, T.H. 1888. Report on the fishes observed in Great Egg Harbor Bay, New Jersey, during 

the summer of 1887. Bulletin of the United States Fisheries Commission 7: 129-154. 

Bearden, C.M. 1960. Flounders and their cousins - unique fish. South Carolina Wildlife. 5 pp. 

. 1961. Common marine fishes of South Carolina. Contribution Bears Bluff Laboratory 

34. 47 pp. 

Bearden, C.M. and C.H. Farmer, 111. 1972. Fishery resources of Port Royal Sound estuary. 

Pages 204-212 in Port Royal Sound environmental study. South Carolina Water Resources 

Commission. Columbia. 

Bearden, C.M., R. Low, R. Rhodes, R. Van Dolah, C. Wenner, E. Wenner and D. Whitaker. 1985. 

A review and analysis of commercial shrimp trawling in the sounds and bays of South Carolina. 

Technical Report No. 61. South Carolina Marine Resources Center. 51 pp. 

Beatty, H.R., R.P. Webster and E.L. Wenner. 1989. Temporal and spatial variations in biomass, 

abundance and community composition of fishes and invertebrates from the coastal habitat, 

South Atlantic Bight, 1987- 1988. SeamapSA Annual Report. South Carolina Marine Resources 

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Bedsole, H.L., Jr., B.F. Holland, Jr. and J.W. Gillikin, Jr. 1980. State of North Carolina RIV Dan 

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Berg, D.L. and J.S. Levinton. 1985. The biology of the Hudson-Raritan estuary, with emphasis 

on fishes. NOAA Technical Memorandum NOS OMA 16. National Oceanic and Atmospheric 

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Berg, M.B. and P. Weller. 1967. Studies of organic acid, sugar, and amino acid transport in 

isolated perfused flounder tubules. Bulletin of the Mount Desert Island Biological Laboratory 7: 4- Z 

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Thomas and Associates, editor. Ecological studies in the bays and waterways near Little Egg 

Inlet and in the ocean in the vicinity of the proposed site for the Atlantic Generating Station, New 

Jersey. Progress Report for January - December 1975. Ichthyological Associates Incorporated, 

Absecon, N.J. 
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Bodammer, J.E. 1980. Ultrastructural studies on the developing cranial cartilage in several 

species of larval-stage fish. International Council for the Exploration of the Sea Council Meeting 

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Bolz, G.R., R.G. Lough and D.C. Potter. 1981. Autumn and winter abundance and distribution of 

ichthyoplankton on Georges Bank and Nantucket Shoals, 1974-76, with special emphasis on 

dominant species. Rapports et Proces-Verbaux des Reunions, Conseil Permanent International 

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stock assessment program bottom trawl survey annual data summary report series, Volume 1990. 

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Bozeman, E.L., Jr. and J.M. Dean. 1980. The abundance of estuarine larval and juvenile fish in a 

South Carolina intertidal creek. Estuaries 3(2): 89-97. 

Breder, C.M., Jr. 1922. The fishes of Sandy Hook Bay. Zoologica 2(15): 331-351. 

. 1948. Field book of marine fishes of the Atlantic coast. G.P. Putnam and Sons, New 

York. 322 pp. 

Briggs, J.C. 1958. A list of Florida fishes and their distribution. Bulletin of the Florida State 

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Briggs, P.T. 1962. The sport fisheries of Great South Bay and vicinity. New York Fish and 

Game Journal 9(1): 1-36. 

. 1965. The sport fishery in the surf on the south shore of Long Island from Jones Inlet to 

Shinnecock Inlet. New York Fish and Game Journal 12(1): 31-47. 

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-- . 1984. Fish investigation in the vicinity of a sewer outfall under construction off the south 

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59. Bullis, H.R. and J.R. Thompson. 1965. Collections by the exploratory fishing vessels Oregon, 

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63. Burke, J.S., J.S. Miller and D.E. Hoss. 1991. Immigration and settlement pattern of Paralichthys 

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64. Burns, R.W. 1974. Seasonal abundance and diversity of larval fishes in a high-marsh tidal 

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65. Burreson, E.M. 1981. Effects of mortality caused by the hemoflagellate Trypanoplasma bullocki 

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504. 
Author index to the Bibliography 

Note: Numbers correspond to citations listed in Appendix A 

Abbe, G.R., 1 

Able,K.W., 2,282,293,284,516,517,518,603, 

604, 605, 664 

Adams, S.M., 3, 4, 608 

Agnello, R.J., 5 

Ahlstrom, E.H., 6, 224 

Allen,D.M., 7,8,9, 10,449 

Almeida, F.P., 11 

Alperin, I.M., 12 

Ames, W.H., 13 

Arnoaka, K., 6 

Anderson, E.D., 14 

Anderson, V.T. Jr., 15 

Anderson, W.W., 16, 17 

Angelovic, J.W., 47 1 

Arnold, E.L., 18 

Arve,J., 19 

Atlantic States Marine Fisheries Commission 

Subcommittee for Fisheries of 

the Atlantic Bight, 20 

Atlantic States Marine Fisheries Commission 

Summer Flounder 

Subcommittee, 2 1, 22 

Atran, S., 672 

Ayvazian, S.G., 126 

Azarovitz, T.R., 23, 24, 74, 208, 653 

Bailey, R.M., 26, 503, 504 

Baker, B.M., 649 

Baker, S., 147 

Baptist, J.P., 27 

Barans, C.A., 635, 636 

Barker, D.L., 7 

Bass, D.G. Jr., 28 

Bayless, J., 291 

Bean, B.A., 61 1 

Bean, T.H., 29 

Bearden, C.M., 30, 31, 32, 33 

Beatty, H.R., 34, 637 

Bedsole, H.L. Jr., 35 

Beirne, C., 669, 670 

Bengston, D.A., 40 

Berg, D.L., 36 

Berg, M.B., 37 

Berrien, P.L., 97 

Berry, F.H., 635, 636 

Bevacqua, E.S., 23 

Bieder, R.C., 38 

Bigelow, H.B., 39 

Birdsong, R.S., 107 

Bisbal, G.A., 40 

Bodammer, J.E., 41, 31 1 

Boehlert, G.W., 452 

Bolz, G.R., 42 

Bond, C.E., 503, 504 

Bonzek, C.F., 43,44, 190 

Bowman, E., 209 

Bowman, R.E., 1.5 1, 302, 303, 347 

Bozeman, E.L. Jr., 45 

Breder, C.M. Jr., 46, 47, 436 

Briggs, J.C., 48 

Briggs, P.T., 49, 50, 51, 52 

Brooker, J.R., 503, 504 

Brooks, H.A., 630 

Brown, B.E., 88, 89, 650 

Brown, C., 506, 507 

Brown, R.A., 53 

Brown, J.T., 220 

Bruce, R.A., 54 

Buchanan, C.C., 55, 461 

Buckley, L.J., 56, 57 

Buller, R.J., 58 

Bullis, H.R., 59 

Bulloch, D.K., 60 

Burke, J.S., 61, 62, 63, 376, 624 

Burnett, J.M., 11, 446 

Burns, R.W., 64 

Burreson, E.M., 65, 66, 67, 68, 69, 70, 71, 72, 

263, 602 

Bushing, M.F., 73 

Byrne, C.J., 14, 23, 24, 74 

Byron, R.R., 96 

Cable, L.E., 231, 732 
Cable, R.M., 75 

Cain, R.E., 76 

Cali, A., 306 

Carley, D.H., 78 

Carlson, J.K., 79 

Carolina Power and Light Co., 80 

Casey, J.F., 81 

Castagna, M., 501 

Castaneda, RE., 11 

Chadwick, D.L., 108, 110 

Chang, S., 82, 83 

Chester, A.J., 229 

Christensen, D.J., 84, 85, 92, 185 

Clark, J.R., 86, 87, 143 

Clark, S.H., 88, 89 

Clayton, G., 90 

Clements, L.C., 91 

Clifford, W.J., 84,85,92 

Clymer, J.P. 111, 8 

Coates, P.G., 172 

Coduri, R.J., 93 

Cole, C., 90 

Cole, R.W., 94, 95, 

Colton, J.B. Jr., 96, 97, 98, 334, 335, 336 

Colton, M.D., 99 

Colvocoresses, J.A., 43, 44, 100, 190 

Connecticut Division of Marine Fisheries, 101, 

102, 103 

Conover, D.O., 363 

Cooke, D.W., I99 

Coon, W.P. 11, 639, 640 

Copeland, B.J., 104, 105, 227 

Coston, L.C., 246, 247 

Coston-Clements, LC., 425, 530, 53 1 

Cowan, J.H. Jr., 107 

Curley, J.R., 108, 109, 110, 17 1 

Dahlberg, M.D., 111, 112, 113, 114, 115, 

Daiber, F.C., 116, 117, 118, 119, 120, 573 

Daniel, L.B. Ill, 121, 638 

Davidson, L., 540, 541 

Davis, R.W., 631 

Dawson, C.E., 122, 123 

Deacutis, C.F., 124 

Dean, J.M., 45, 77, 553 

Decker, E.A., 125 

Deegan, L.A., 126 

DeGroot, S.J., 127, 128, 129 

Delaney, G.R., 130, 175 

Derickson, W.K., 13 1 

Dery, L.M., 132, 133, 466, 574 

Desfosse, J.C., 134, 125 

Despres, L.I., 23 

Despres-Patanjo, L.I., 24, 135, 136, 672 

de Sylva, D.P., 137, 138, 545 

Deubler, E.E. Jr., 140, 141, 662, 663 

Deuel, D.G., 142, 143, 323 

DeVries, D.A., 144, 145, 513 

Dickinson, C.L., 427 

Dillmann, D.W., 57 

Dipiero, S.J., 564 

Donohoe, C.J., 199 

Douglas, J., 505 

Dovel, W.L., 146 

Draxler, S., 593 

Drewry, G.E., 342 

Dudley, D.L., 606 

Duncan, J., 540, 541 

DuPaul, W., 147 

Easley, J.E. Jr., 106 

Ecological Analysts Engineering Science and 

Technology, Inc., 148 

Ecologcal Analysts, Inc., 149 

Edwards, R.L., 150, 151, 152 

Eerly, S.P., 153, 154, 515 

Elsenberg, M., 159 

Eklund, A., 155 

Eldridge, P.J., 156 

Esslg, R.J., 157 

Estes, A.D., 134, 135 

Erickson, M.C., 125 

Estes, A.D., 406 

Estrella, B.T., 249 

Evermann, B.W., 275 

Fahay, M.P., 2, 87, 97, 158, 386, 387, 580 

Fahy, W.E., 140 

Farmanfarmaian, A., 159 

Farmer, C.H. 111, 32 

Fee, R., 160 

Feigenbaum, D.L., 73 

Festa, P.J., 161, 162, 163, 164, 165, 166, 167, 

168, 396, 397, 398 

Figley, W., 169, 170 

Finn, J.T., 126 

Fiske, J.D., 109, 17 1, 172 

Fitch, J.E., 26 

Fitzhugh, G.R., 376 

Florida Department of Natural Resources, 173 

Fogarty, M.J., 174. 175 

Foster, D.B., 335, 336 

Foster, H.A., 23, 24 
Fowler, H.W., 176, 177, 178, 179, 180, 181, 182, 

183, 551, 61 1 

Freeman, B.L., 184, 185, 186 

Frisbie, C.M., 78 

Frizzell, L.J., 68 

Gabriel, W.L., 187 

Gahn, T., 594, 595, 596, 598 

Gaichas, S.K., 270 

Gallagher, M.L., I88 

Gartner, J.V., 189 

Geer, P.J., 43, 44, 190 

Gehringer, J.W., 17 

Gerhart, E.H., 191 

Gerry, L.R., 632 

Gibson, C.I., 355 

Gilbert, C.R., 192 

Gillikin, J.W. Jr., 35, 175, 193, 194, 195, 196, 482, 

483 

Gilmore, R.G. Jr., 197. 198, 199, 274 

Ginsburg, I., 200 

Ginter, J.J.C., 364 

Goldman, J.C., 201 

Goldstein, R.J., 202 

Goode, G.B., 203 

Gordon, B.L., 204 

Goss, D.K., 276 

Grant, G.C., 458, 459 

Graunke, B., 274 

Greenfield, R.C., 11 

Greenwood, P.H., 205 

Greges, M.P., 564 

Grimes, B.H., 206 

Grosslein, M.D., 25, 207, 208, 209, 210 

Gudger, E.W., 211, 212, 214 

Gunter, G., 214 

Gusey, W.F., 215 

Gutherz, E.J., 216 

Guthrie, R.O., 195 

Hackney, C.T., 520 

Haire, M.S., 312 

Halgren, B., 217 

Hall, W.M. Jr., 670 

Hamer, P.E., 152, 218, 314 

Hardy, J.D. Jr., 333 

Hargis, W.J. Jr., 219 

Harrell, M.L., 188 

Harris, C.D., 324, 325, 326, 327 

Harris, R.E., 43, 44, 190 

Harvell, C.H., 145 

Hassler, W.W., 220 

Hattala, K., 672 

Hawkes, J.W., 239 

Hawkins, J.H., 221, 513 

Heck, K.L. Jr., 453 

Henderson, E.M., 22 

Hennemuth, R.C., 223 

Hensley, D.A., 6, 224 

Herald, E.A., 26 

Herman, S.S., 8, 225, 286 

Herold, R.C., 226 

Herrema, D.J., 199 

Hester, J.J., 227 

Hettler, W.F. Jr., 228, 229, 247, 609 

Heyerdahl, E.G., 210 

Hickey, J.M., 108, 109 

Hicks, B., 230 

Hildebrand, S.F., 23 I, 232, 233 

Hill, N.L., 220 

Hillman, R.J., 234 

Himchak, P.J., 235, 3-36, 237, 238 

Hocutt, C.H., 269, 270 

Hodgins, H.O., 239 

Hodson, R.G., 104, 105, 106, 632 

Hoese, H.D., 240 

Hoff, F.H. Jr., 610 

Hoff, J.G., 575 

Hoffman, W.G. 11, 241 

Hogarth, W.T., 539 

Holder, D.R., 248 

Holland, A.F., 3 12 

Holland, B.F. Jr., 35, 195, 196, 242, 482, 483 

Holliday, M.C., 157 

Horwitz, R.J., 243 

Hoss, D.E., 62, 63, 91, 244, 245, 246, 247, 609, 

646 

Hottell, H.E., 248 

Howe, A.B., 249, 672 

Howell, W.H., 77 1, 272 

Hsia, R., 159 

Hughes, E.H., 250, 251 

Huish, M.T., 206 

Hultin, H.O., 125 

Hunninen, A.V., 75 

Hussakoff, L., 252 

lannaccone, V., 159 

Ichthyological Associates, 254, 255, 256, 257, 

258 

Industrial Fishery Project, 259 

International Gamefish Association, 260 

Irlandr, L.R., 425 
Ishibashi, N., 261 

Itzkowitz, N., 262, 564 

Jacobs, F., 312 

Jansen, M.E., 263 

Jearld, A. Jr., 574 

Jeffries, H.P., 264, 265 

Jensen, A.C., 266, 267, 268 

Jensen, J., 312 

Jesien, R.V., l I, 269, 270 

Johns, D.M., 27 1, 272 

Johnson, G.N., 612, 613, 614 

Johnson, K.L., 273 

Johnson, W.C., 264 

Jones, R.S., 274 

Jordan, D.S., 275, 276 

Jorgenson, S.C., 277, 375 

Joseph, E.B., 344, 345, 552 

Judy, M.H., 278 

June, F.E., 279 

Junqueira, L.C., 280, 281 

Kalber, F.A., 137, 138 

Kasper, V., 594, 595, 596, 598 

Keefe, M., 282, 283, 284 

Keirans, W.J. Jr., 285, 286 

Keiser, R.K., 287 

Keisler, A.S., 626 

Kemmerle, S., 288 

Kendall, A.W. Jr., 87, 97, 289 

Kerby, J.H., 206 

Kernehan, R.J., 622 

Kerr, G.A., 290 

Keup, L., 291 

Kimmel, J.J., 292 

Kindred, J.E., 1971 

Kirsch, R., 294 

Kjelson, M.A., 472, 609 

Klein-MacPhee, G., 272, 295, 296, 297 

Krouse, C.W., 530, 53 1 

Kulczyck, G.R., 274 

Kuntz, A., 298 

Kyle, H.M., 299 

Lachner, E.A., 26, 503, 504 

LaCroix, M.W., 608, 609 

Lange, A.M.T., 14, 300 

Langton, R.W., 301, 302, 303, 

Lascara, V.J., 304 

Latham, R., 305 

Laudan, R., 306 

Laurence, G.C., 307 

Lawton, R.P., 108, 109, 110, 171 

Leim, A.H., 308 

Lesser, C.A., 309 

Levinton, J.S., 36 

Lewis, E.J., 3 10, 3 11 

Lewis, R.R., 562 

Lindsey, C.C., 26 

Lipcon, V., 598 

Lison, A.J., 3 12, 3 13, 

Livingstone, R., 152 

Livingston, R.J., 556 

Lockwood, K., 170 

Longo, J., 564 

Lough, R.G., 42 

Low, R., 33 

Lucy, J.A., 341 

Lugger, O., 616 

Lux, F.E., 218, 314, 315, 316, 317, 318, 319 

Lynch, T.R., 320 

Lyons, P., I35 

MacKenzie, C.L. Jr., 321 

MacPhee, G., 322 

Maddox, M.B., 638 

Magley, W.C., 274 

Mahoney, J.B., 315, 323, 561 

Mahood, R.K., 324, 325, 326, 327 

Malins, D.C., 618 

Malloy, K.D., 328, 329, 330, 331 

Malsberger, R.G., 286 

Mann, W.C., 310 

Manooch, C.S. 111, 332 

Mansueti, A.J., 333 

Marak, R.R., 334, 335, 336 

Marcellus, K.L., 337 

Marsh, E., 338 

Marshall, A., 339, 340, 341 

Martin, F.D., 342 

Martore, R.M., 583, 637 

Mason, D., 540, 541 

Mason, J.M., 14 

Massmann, W.H., 343, 344, 345,490 

Mast, S.O., 346 

Matheson, RE., 2 

Maurer, R.O. Jr., 347 

Mayo, R.K., 348, 349, 446, 562 

McAdams, M., 540, 541 

McAllister, D.E., 350 

McClain, J.F. Jr., 351, 352 

McDermott, V., 353, 354 

McEachran, J.D., 407 

McErlean, A.J., 355 
McGovern, J.C., 356, 357, 637 

McHugh, J.L., 358, 359, 360, 361, 362, 363, 364 

McKenzie, R.A., 620 

McKown, K.A., 672 

McLaughlin, S.M., 3 1 1 

McMillan, D.G., 582, 657 

Mercer, L.P., 365 

Meredith, A., 288 

Merriman, D., 366, 623 

Metzger, F. Jr., 367 

Michels, S.F., 368 

Michener, W.K., 9, 10, 449 

Mid-Atlantic Council Staff, 369 

Mid-Atlantic Fishery Management Council, 370, 

371, 372, 373, 374 

Midlige, F.H., 323 

Miglanese, J.V., 552 

Mihursky, J.A., 355 

Miller, D., 336, 590 

Miller, G.L., 277, 375 

Miller, J.M., 376, 377, 378 

Miller, J.S., 63 

Miller, R.W., 379 

Monaco, M.E., 425 

Monaghan, J.P. Jr., 365, 380a, 380b, 491 

Monroe, R.J., 104, 437, 438 

Moore, H.F., 381 

Moran, D., 306 

Moran, J.E., 638 

Moran, R.L., 313 

Moran-Johnson, R.L., 312 

Morgan, M.A., 564 

Morris, T.L. Jr., 382 

Morse, W.W., 2, 383, 384, 385, 386, 387, 651, 

652, 653, 654, 655, 656 

Moser, H.G., 6 

Murawski, S., 90, 388, 389, 390, 391, 392, 393, 

394, 395, 396, 397, 398, 399 

Murchelano, R.A., 136, 400, 401, 507, 523 

Music, J.L. Jr., 324, 325, 326, 327, 402, 403 

Musick, J.A., 100, 134, 135, 404, 405, 406, 407 

Myer, G.S., 205 

Myrvik, Q.N., 626 

Nagle, J.J., 593, 594, 595, 596, 598 

National Marine Fisheries Service, 408-423 

Nelson, D.M., 425 

Nesbit, R.A., 426 

Neville, W.C., 426, 427, 642 

New Jersey Department of Environmental 

Protection, 428-433 

Newman, M.W., 434 

Nichols, J.T., 435, 436 

Nicholson, N., 460 

Nichy, F.E., 316, 317, 318 

Nixon, S.W., 456 

Noble, E.B., 437, 438 

Norcross, J.J., 344, 345 

Norman, J.R., 439 

North Carolina Division of Marine Fisheries, 

440-444 

Northeast Utilities Service Company, 445 

O'Brien, L., 446 

O'Conner, J.S., 447, 448 

O'Connor, S.G., 355 

Odum, E.P., 115 

Ogburn, M.V., 449 

Olla, B.L., 450 

Olney, J.E., 45 1, 452 

O'Neil, S.P., 575 

Orth, R.J., 453 

Osborn, C.M., 454, 455 

Oviatt, C.A., 456 

Pacheco, A.L., 83, 457, 458, 459, 657 

Pafford, J.M., 402, 460 

Palmer, B.A., 324, 325, 326, 327 

Pancirov, R.J., 53 

Parker, L.L., 73 

Parker, R.O., 461 

Parrish, J., 90 

Pearcy, W.G., 462 

Pearson, J.C., 463, 464 

Pendleton, E.C., 465 

Pentilla, A., 466 

Perlmutter, A., 467, 468 

Perra, P., 469 

Perschbacher, P., 540, 541 

Peters, D.S., 377, 470, 47 1, 

Phalen, P.S., 473 

Phelan, B.A., 657 

Pietrafesa, L.J., 378 

Pikanowski, R.A., 657 

Podgar, T.T., 3 12 

Poole, J.C., 12, 175, 314, 474, 475, 476, 477, 

478,479 

Porter, H.J., 542 

Porter, L.R. Jr., 319 

Potter, D.C., 42 

Powell, A.B., 480, 481, 482, 483, 484, 485, 486 

Powles, H., 487 

Price, K. Jr., 131 
Price, K.S., 545 

Price, T.J., 27 

Pritchard, E.S., 24 

Pulley, M.G., 488 

Purvis, C., 489 

Radcliffe, L., 298 

Ralph, D.E., 175, 652, 653, 654, 655, 660, 672 

Rand, H.G. Jr., 93 

Raney, E.C., 490 

Ray, G.C., 505 

Raynie, R.C., 81 

Reback, K., 108, 1 10 

Reed, J.P., 378 

Reed, L., 491 

Reichert, M.J.M., 392 

Reinach, F., 280 

Reintjes, J.W., 279, 493 

Reis, R., 494 

Reisersen, L., 495 

Rhode Island Division of Fish and Wildlife, 496 

Rhodes, R., 33 

Rice, C.D., 627 

Richards, A., 497 

Richards, C.E., 498, 499, 500, 501 

Richards, S.W., 462, 502 

Richkus, W.A., 312 

Riggs, S.R., 105, 106 

Robins, C.R., 503, 504, 505 

Robohm, R.A., 506, 507, 508 

Rogers, S.G., 403, 509, 5 I 0 

Roithmayr, C.M., 493 

Rosen, D.E., 205 

Rosenberg, M., 511 

Ross, J.L., 365, 491, 512, 513 

ROSS, S.W., 154, 5 14, 5 15 

Roumillat, W.A., 637, 638 

Rountree, R.A., 516, 517, 518, 605 

Rozas, L.P., 519, 520 

Rulifson, R.A., 188, 521, 522 

Ryder, C., I26 

Salles, L.M.M., 280, 281 

Samet, C.E., 450 

Sandifer, P.S., 639, 630, 64 1 

Sandoy, K., 540, 54 1 

Sass, S.L., 523 

Sawyer, R.T., 524 

Sawyer, T.K., 3 1 1 

Scarlett, P.G., 175, 2 17, 525, 526, 527, 528, 574 

Scattergood, L.W., 529 

Schaaf, W.E., 246, 530, 53 1 

Schaefer, R.H., 532 

Schauss, R.P., Jr., 533 

Schroeder, W.C., 39, 233, 534 

Schubel, J.R., 262, 564 

Schwartz, F.J., 484, 485,486, 535, 536, 537, 538, 

539, 540, 541, 543- 

Sclar, R.C., 366 

Scott, W.B., 26, 308, 543 

Scott, R.N., 503, 504 

Scotton, L.N., 544, 545 

Seagraves, R.J., 94, 95, 546, 547, 548, 549 

Sedberry, G.R., 634 

Settle, M.E., 425 

Sharp, B., 551 

Sharp, J.H., 550 

Shealy, M.H. Jr., 552, 639, 640, 641 

Shenker, J.M., 553 

Shepherd, G.P., 2, 85, 554, 555 

Sheridan, P.F., 556 

Sherman, K., 557, 558 

Sherr, E.B., 251 

Shipman, S., 559 

Shuster, C.N. Jr., 138 

Shuster, C.W., 560 

Sibunka, J.D., 581, 660 

Sicluna, J.D., 672 

Silverman, M.J., 561, 658, 659 

Sinderman, C.J., 601 

Sissenwine, M.P., 562 

Smigielski, A.S., 563 

Sm~th, C.F., 564 

Smith, H.M., 565, 566, 567 

Smith, J.W., 568, 569, 638 

Smith, M.G., 543 

Smith, N.S., 545 

Smith, R.E., 545 

Smith, R.L., 570 

Smith, R.W., 120, 175, 571, 572, 573, 574, 

Smith, S.I., 619 

Smith, S.M., 575 

Smith, W.G., 87, 97, 386, 387, 558, 576-582, 660 

Socci, R., 159 

Sparrow, D.S., 508 

Spear, H.S., 58 

Spitsbergen, D., 582 

Stancyk, S.E., 9, 10 

Steady, E.J., 653, 655 

Stehlik, L.L., 656, 657 

Stender, B.W.. 583, 635, 636 

St. Onge, J.M., 98 
Biblioaraphv 

Stender, B.W., 487 

Stephan, C.D., 584, 585 

Stephan, D.C., 473 

Stern, H., Jr., 210 

Stickney, R.R., 586, 587, 588, 589, 590, 644, 645 

Stock Assessment Workshop, 59 1, 592 

Stolen, J.S., 306, 593, 594, 595, 596, 597, 598, 

Stone, R.B., 461 

Struhsaker, P., 599 

Studholme, A.L., 450 

Sumida, B.Y., 6 

Sumner, F.B., 600 

Swanson, R.L., 601 

Sypek, J.P., 69, 602 

Szedlmayer, S.T., 603, 604, 605 

Tagatz, M.E., 606 

Targett, T.E., 155, 329, 330, 331, 509 

Tate, J., 541 

Temple, R.F., 99 

Terceiro, M., 265, 555, 607 

Thayer, G.W., 608, 609 

Thompson, J.R., 59 

Tinsman, J.C., 94, 95 

Topp, R.W., 610 

Truitt, R.V., 61 1 

Turner, W.R., 612, 613, 614 

Tyler, A.V., 615 

Uhler, P.R., 616 

Van Den Avyle, M.J., 510 

Van Ber Veer, H.W., 492 

Van Dolah, R., 33 

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., 62 1 

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 

Weinstein, M.P., 539, 575, 628, 629, 630, 631, 

632, 633 

Weiss, S.L., 632, 633 

Weitzman, S.H., 205 

Weller, P., 37 

Wells, A., 581, 582 

Wenner, C.A., 33, 357, 569, 634, 635, 636, 637, 

638 

Wenner, E.L., 33, 34, 639, 640, 641 

Wesche, A.E., 81 

Westman, J.R., 427, 642 

Wheatland, S.B., 043 

White, D.B., 587, 588, 589, 590, 644, 645 

White, J.C. Jr., 141, 646 

White, R.L., 399 

Widerstrom, F.L. Jr., 647 

Wilk, S.J., 175, 648-660 

Williams, B.C., 403 

Williams, A.B., 661, 662, 663 

Witting, D.A., 664 

Witzig, J.F., 157 

Wolff, M., 665, 666, 667 

Woodward, A.G., 668 

Woolcott, W.S., 669, 670 

Wyanski, D.M., 67 1 

Young, B.H., 672 

Younger, R.R., 673 

Zamos, J.A., 673 

Zingmark, R.G., 671 

Ziskowski, J.J., 400, 401, 672 

Zwerner, D.E., 70, 71, 72 

Subject index to the Bibliography 

Note: Numbers correspond to citations listed in Appendix A 

adults: 144 

age: 11, 132, 133,466,474,48 1, 554 

anatomy: 128, 129, 226, 280, 281, 293, 294, 299, 338, 382, 455, 

669, 670 (see also anomalies) 

anomalies: 15, 40, 11 1, 123, 140, 189, 21 1, 212, 213, 252, 484, 589, 646 

aquaculture: 587, 588, 589, 644 

Baltimore Canyon: 53 

behavior: 127, 295, 296, 304, 346, 450, 590 

bibliography: 123, 495, 526, 545, 645 

biochemistry: 37, 125, 159, 188, 330, 33 1, 5 11 

biomass: 88, 3 19, 5 16, 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, 120, 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 

diseaseiparasites: 65, 66, 67, 68, 69, 70, 71, 72, 75, 122, 136, 219, 239, 

263, 306, 31 1, 323, 400, 401, 434, 506, 507, 508, 523, 524, 561, 588, 593, 594, 595, 596, 597, 598, 602, 

67 2 

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: 1 19, 534, 586 

early life history: 2, 158, 376, 458, 510, 622 
eggs: 2, 42, 96, 98, 146, 225, 229, 237, 241, 271, 272, 278, 285, 286, 298, 333, 334, 335, 336, 352, 354, 366, 

451,452, 457, 467, 487, 557, 576, 577, 580, 582, 621, 637, 643 

electrophoresis: 93, 6 17 

entrainment: see industryienergy plant studies 

estuarine dependence: 87, 378 

fecundity: see reproduction 

feeding: 61, 128, 129, 261, 292, 304, 328, 329, 330, 382, 450, 477, 590 

fisheries (commercialirecreational): 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, 2 10, 235, 249, 259, 260, 266, 267, 268, 273, 279, 287, 288, 300, 309, 3 16, 317, 320, 

324, 325, 326, 327, 339, 340, 341, 351, 359, 360, 361, 362, 363, 364, 365, 369, 370, 371, 372, 373, 374, 

379, 394, 398,402, 403, 408, 409, 410, 41 1,412, 413,414,415,417,418,419, 421, 422, 423, 426, 438, 

44 1, 442, 443, 460, 463, 468, 469, 473, 475, 476, 49 1, 493, 496, 498, 499, 5 12, 5 13, 5 14, 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 

foodlfood 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, 11 1, 112, I 13, 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, 328, 329, 330, 331, 470, 47 1, 475, 481, 492, 5 17, 574, 603, 605 

habitat selection/use: 61, 62, 282, 322, 437, 510, 520, 534, 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 industryienergy plant studies 

industryienergy 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, 238, 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. lengthiweight): 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, 53 I, 621, 638, 642, 654 

Long Island Sound: 427, 445, 502 

marking: see tagging~rnarking 

Maryland: 19, 81. 179,369,270, 312, 313, 535, 536, 537, 611, 616 
Massachusetts: 90, 108, 109, 110, 126, 172, 249, 317, 523, 551, 566, 576, 581 

maturity: see reproduction 

mesh selectivity: 14, 194, 195, 217, 668 

meristcs: 130 

metabolism: see physiology 

metamorphosis: 283, 299 

Middle Atlantc: 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 

mortalityisurvival: 65, 72, 297, 328, 329, 330, 492, 601, 605, 625 

movement: 145, 193, 218, 242, 245, 269, 270, 305, 378, 380a, 604 (see also taggingimarking) 

Nantucket Shoals: 42, 152, 186 

New England: 18, 150, 266, 3 18, 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, 

18 1, 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, 41 1, 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, 515, 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: 9 1, 184, 550 

parasites: see diseaseiparasites 

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, 21 1, 213, 252, 280, 281, 346, 454, 455, 589, 600, 646 

pollutantsipoIlution: 9, 27, 53, 82, 112, 159, 190, 201, 239, 244, 246, 447, 448, 507, 508, 530, 531, 595, 597, 

598, 618, 657 
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, 385, 397, 399, 424, 446, 563, 651, 656, 579 

Rhode Island: 204, 225, 264, 265, 320, 456, 496 

salinity: 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, 5 19, 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, 64 1, 67 1 

spawning: see reproduction 

species list: 26, 176, 177, 183, 197, 214, 350, 352, 377, 381, 404, 490, 503, 504, 61 1, 616, 671 

survival: see mortality/survival 

stocks/stock assessment: 8 1, 130, 134, 135, 144, 175, 207, 22 1, 269, 270, 406, 497, 607, 617 

systematics: see identification 

tagginglmarking: 145, 193, 218, 242, 245, 269, 270, 318, 380, 393, 406 

temperature: 134, 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, 453, 608, 630 

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. lengthiweight) 
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., NOAAiNational 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 02881 

Burke, John S., NOAAiNational Marine Fisheries Service, Southeast Fisheries Science Center, Beaufort 

Laboratory, 101 Pivers Island Rd., Beaufort, NC 285169722 

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 Office Bldg., Annapolis, MD 21401 

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

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

Daniel, Louis B., Ill Virginia Institute of Marine Science, College of William and Mary, P.O. Box 786, 

Gloucester Point, VA 23062 

Desfosse, Joseph C., Virginia lnstitute of Marine Science, College of William and Mary, P.O. Box 1046, 

Gloucester Point, VA 23062 

DeVries, Doug, NOAAiNational 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., NOANNational Marine Fisheries Service, Northeast Fisheries Science Center, 

James J. Howard Marine Sciences Laboratory, Highlands, NJ 07732 

Fogarty, Michael NOAAiNational Marine Fisheries Service, Northeast Fisheries Science Center, Woods 

Hole Laboratory, 116 Water Street, Woods Hole MA 07543-1097 

Fonseca, Mark NOANNational Marine Fisheries Service, Southeast Fisheries Science Center, Beaufort 

Laboratory, 10 1 Pivers Island Rd., Beaufort, NC 285 16-97?? 
B-2 

Experts 

Gorski, Stan, NOAAiNational Marine Fisheries Service, Northeast Fisheries Science Center, James J. 

Howard Marine Sciences Laboratory, Highlands NJ 07732 

Hettler, William, NOAAiNational Marine Fisheries Service, Southeast Fisheries Science Center, Beaufort 

Laboratory, 101 Pivers Island Rd., Beaufort, NC 285 16-9722 

Hocutt, Charles H., Horn Point Environmental Laboratory, University of Maryland, P.O. Box 775, 

Cambridge, MD 2 16 13-0775 

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. NOAAiNational 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., lnstitute of Marine and Coastal Sciences, Rutgers University, Marine Field Station, 

800 Great Bay Blvd., Tuckerton, NJ 08087 

Keefe, MaryLouise, Oregon Department of Fish & Wildlife, 21 1 lnlow Hall, Eastern Oregon State 

College, LaGrande, OR 97850 

Klein-MacPhee, Grace, University of Rhode Island, Graduate School of Oceanography, South Ferry 

Road, Narragansett, RI 02887- 1 197 

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, 89 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.0 Box 358, 

Gloucester Point. VA 23062 
Experts 

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 367, Melvin Village, NH 03850 

Powell, Allyn B., NOAAiNational Marine Fisheries Service, Southeast Fisheries Science Center, 

Beaufort Laboratory, 101 Pivers lsland Rd., Beaufort, NC 285169722 

Powell, Christopher, Rhode lsland 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., NOAAiNational Marine Fisheries Service, Northeast Fisheries Science Center, 

Woods Hole Laboratory, 1 I6 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., NOAAiNational 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., 1 Conservation 

Way, Brunswick, GA 3 1523-8600. 

Smith, Joseph W., NOANNational 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 1431, Dover, DE 

19903 

Smith, Walter G., NOANNational Marine Fisheries Service, Northeast Fisheries Science Center, 

James J. Howard Marine Sciences Laboratory, Highlands NJ 07732 

Studholme, Anne L., NOAAiNational 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, NOAAiNational Marine Fisheries Service, Northeast Fisheries Science Center, Woods 

Hole Laboratory, 116 Water Street, Woods Hole, MA 02543-1097 

Wenner, Charles A. South Carolina Wildljfe and Marine Resources, P.O. Box 12559, Charleston, SC 

29412-2559 

Wilk, Stuart J., NOAAiNational Marine Fisheries Service, Northeast Fisheries Science Center, James J. 

Howard Marine Sciences Laboratory, Highlands NJ 07732 
8-4 

Experts 

Wyanski, David M., South Carolina Wildlife and Marine Resources, P.O. Box 12559, Charleston, SC 

294 1 2 
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 

5600 Old Dixie Highway 

Ft. Pierce, FL 33946 

Attention: R.G. Gilmore, JI 

Institute, Inc. 

Georgia Department of Natural Resources 

Coastal Fisheries Dept., 1 Consewation 

Way, Brunswick, GA 3 1523-8600. 

Attention: J.L. Music, Jr., J. Pafford, S. Shipman 

Maryland Department of Natural Resources 

Tidewater Administration 

Fisheries Division, 580 Taylor Avenue 

Tawes State Off ice Bldg. 

Annapolis, MD 21401 

Attention: J.F. Casey 

Massachusetts Division of Marine Fisheries 

Cat Cove Laboratory, Salem, MA 01970 

Attention: W. Castonguay 

NOAAiNational Marine Fisheries Service 

Southeast Fisheries Science Center 

Beaufort Laboratory 

101 Pivers Island Road 

Beaufort, NC 285 16 

Attention: J. Burke, S.E. Chester, L. Coston-Clements, D. Hoss, D. Peters, A.B. Powell 
User aroups 

NOAAiNational 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 

NOAAiNational 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 1 1790 

Attention: K. McKown, V.J. Vecchio, B.H. Young 

State University of New York 

Marine Science Research Center 

Stony Brook, NY 1 1794-5000 

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 qrouos 

North Carolina Division of Marine Fisheries 

Washington District Office, Washington, NC 27889 

Attention: D. 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, R. 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 
User aroups 

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 PRINTING OFFICE: 1995-404-445/25423 


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 ters and larvae hatch. Larvae are pelagic and begin 

development in the ocean It 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.