U. S. DEPARTMENT OF COMMERCE

National Oceanic and Atmospheric Administration

National Marine Fisheries Service

Northeast Region

Northeast Fisheries Science Center

Woods Hole, Massachusetts

November 2000

NOAA Technical Memorandum NMFS-NE-161

Demersal Fish and American Lobster

Diets in the

Lower Hudson - Raritan Estuary
149. Essential Fish Habitat Source Document: Scup, Stenotomus chrysops, Life History and Habitat Characteristics.

By Frank W. Steimle, Christine A. Zetlin, Peter L. Berrien, Donna L. Johnson, and Sukwoo Chang. September 1999.

v + 39 p., 18 figs., 1 table. NTIS Access. No. PB2001-100144.

150. Essential Fish Habitat Source Document: Spiny Dogfish, Squalus acanthias, Life History and Habitat

Characteristics. By Donald G. McMillan and Wallace W. Morse. September 1999. v + 19 p., 10 figs., 3 tables. NTIS

Access. No. PB2001-100148.

151. Essential Fish Habitat Source Document: Summer Flounder, Paralichthys dentatus, Life History and Habitat

Characteristics. By David B. Packer, Sara J. Griesbach, Peter L. Berrien, Christine A. Zetlin, Donna L. Johnson, and

Wallace W. Morse. September 1999. vi + 88 p., 38 figs., 6 tables. NTIS Access. No. PB2001-100149.

152. Essential Fish Habitat Source Document: Tilefish, Lopholatilus chamaeleonticeps, Life History and Habitat

Characteristics. By Frank W. Steimle, Christine A. Zetlin, Peter L. Berrien, Donna L. Johnson, and Sukwoo Chang.

September 1999. v + 30 p., 12 figs., 2 tables. NTIS Access. No. PB2001-100143.

153. U.S. Atlantic and Gulf of Mexico Marine Mammal Stock Assessments -- 1999. By Gordon T. Waring, Debra

L. Palka, Phillip J. Clapham, Steven Swartz, Marjorie C. Rossman, Timothy V.N. Cole, Larry J. Hansen, Kathryn D.

Bisack, Keith D. Mullin, Randall S. Wells, and Neilo B. Barros. October 1999. vii + 196 p., 17 figs., 59 tables, 1 app.

NTIS Access. No. PB2000-102497.

154. Report of the Joint Scientific Review Group Workshop, April 13-14, 1999, Seattle, Washington. By Richard

L. Merrick, compiler. November 1999. v + 22 p., 3 figs., 6 tables, 5 app. NTIS Access. No. PB2000-105393.

155. Food of Northwest Atlantic Fishes and Two Common Species of Squid. By Ray E. Bowman, Charles E.

Stillwell, William L. Michaels, and Marvin D. Grosslein. January 2000. xiv + 138 p., 1 fig., 7 tables, 2 app. NTIS Access.

No. PB2000-106735.

156. Proceedings of the Summer Flounder Aging Workshop, 1-2 February 1999, Woods Hole, Massachusetts. By

George R. Bolz, James Patrick Monaghan, Jr., Kathy L. Lang, Randall W. Gregory, and Jay M. Burnett. May 2000. v

+ 15 p., 5 figs., 5 tables. NTIS Access. No. PB2000-107403.

157. Contaminant Levels in Muscle of Four Species of Recreational Fish from the New York Bight Apex. By Ashok

D. Deshpande, Andrew F.J. Draxler, Vincent S. Zdanowicz, Mary E. Schrock, Anthony J. Paulson, Thomas W. Finneran,

Beth L. Sharack, Kathy Corbo, Linda Arlen, Elizabeth A. Leimburg, Bruce W. Dockum, Robert A. Pikanowski, Brian

May, and Lisa B. Rosman. June 2000. xxii + 99 p., 6 figs., 80 tables, 3 app., glossary. NTIS Access. No. PB2001-107346.

158. A Framework for Monitoring and Assessing Socioeconomics and Governance of Large Marine Ecosystems.

By Jon G. Sutinen, editor, with contributors (listed alphabetically) Patricia Clay, Christopher L. Dyer, Steven F.

Edwards, John Gates, Tom A. Grigalunas, Timothy Hennessey, Lawrence Juda, Andrew W. Kitts, Philip N. Logan, John

J. Poggie, Jr., Barbara Pollard Rountree, Scott R. Steinback, Eric M. Thunberg, Harold F. Upton, and John B. Walden.

August 2000. v + 32 p., 4 figs., 1 table, glossary. NTIS Access. No. PB2001-106847.

159. An Overview and History of the Food Web Dynamics Program of the Northeast Fisheries Science Center,

Woods Hole, Massachusetts. By Jason S. Link and Frank P. Almeida. October 2000. iv + 60 p., 20 figs., 18 tables,

1 app. NTIS Access. No. PB2001-103996.

160. Measuring Technical Efficiency and Capacity in Fisheries by Data Envelopment Analysis Using the General

Algebraic Modeling System (GAMS): A Workbook. By John B. Walden and James E. Kirkley. October 2000. iii

+ 15 p., 9 figs., 5 tables. NTIS Access. No. PB2001-106502.

Recent Issues in This Series:
NOAA Technical Memorandum NMFS-NE-161

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

employ thorough internal scientific review; some issues employ external scientific

review. By design, reviews are transparent collegial reviews, not anonymous peer

reviews. All issues may be cited in formal scientific communications.

U. S. DEPARTMENT OF COMMERCE

Norman Y. Mineta, Secretary

National Oceanic and Atmospheric Administration

D. James Baker, Administrator

National Marine Fisheries Service

Penelope D. Dalton, Assistant Administrator for Fisheries

Northeast Region

Northeast Fisheries Science Center

Woods Hole, Massachusetts

November 2000

Demersal Fish and American Lobster Diets

in the Lower Hudson - Raritan Estuary

Frank W. Steimle, Robert A. Pikanowski, Donald G. McMillan,

Christine A. Zetlin, and Stuart J. Wilk

National Marine Fisheries Serv., 74 Magruder Rd., Highlands, NJ 07732
Editorial Notes

Species Names: The NEFSC Editorial Office’s policy on the use of species names in all technical communications is

generally to follow the American Fisheries Society’s lists of scientific and common names for fishes (i.e., Robins et al.

1991a), mollusks (i.e., Turgeon et al. 1998b), and decapod crustaceans (i.e., Williams et al. 1989c), and to follow the

Society for Marine Mammalogy's guidance on scientific and common names for marine mammals (i.e., Rice 1998d).

Exceptions to this policy occur when there are subsequent compelling revisions in the classifications of species, resulting

in changes in the names of species (e.g., Cooper and Chapleau 1998e

, McEachran and Dunn 1998f).

Statistical Terms: The NEFSC Editorial Office’s policy on the use of statistical terms in all technical communications

is generally to follow the International Standards Organization’s handbook of statistical methods (i.e., ISO 1981g

).

Internet Availability: This issue of the NOAA Technical Memorandum NMFS-NE series is being copublished, i.e., as

both a paper and Web document. The Web document, which will be in HTML (and thus searchable) and PDF formats,

can be accessed at: http://www.nefsc.nmfs.gov/nefsc/publications/.

a

Robins, C.R. (chair); Bailey, R.M.; Bond, C.E.; Brooker, J.R.; Lachner, E.A.; Lea, R.N.; Scott, W.B. 1991. Common and scientific names of fishes

from the United States and Canada. 5th ed. Amer. Fish. Soc. Spec. Publ. 20; 183 p.

b

Turgeon, D.D. (chair); Quinn, J.F., Jr.; Bogan, A.E.; Coan, E.V.; Hochberg, F.G.; Lyons, W.G.; Mikkelsen, P.M.; Neves, R.J.; Roper, C.F.E.;

Rosenberg, G.; Roth, B.; Scheltema, A.; Thompson, F.G.; Vecchione, M.; Williams, J.D. 1998. Common and scientific names of aquatic

invertebrates from the United States and Canada: mollusks. 2nd ed. Amer. Fish. Soc. Spec. Publ. 26; 526 p.

c

Williams, A.B. (chair); Abele, L.G.; Felder, D.L.; Hobbs, H.H., Jr.; Manning, R.B.; McLaughlin, P.A.; Pérez Farfante, I. 1989. Common and

scientific names of aquatic invertebrates from the United States and Canada: decapod crustaceans. Amer. Fish. Soc. Spec. Publ. 17; 77 p.

d

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

e

Cooper, J.A.; Chapleau, F. 1998. Monophyly and interrelationships of the family Pleuronectidae (Pleuronectiformes), with a revised classification.

Fish. Bull. (U.S.) 96:686-726.

f

McEachran, J.D.; Dunn, K.A. 1998. Phylogenetic analysis of skates, a morphologically conservative clade of elasmobranchs (Chondrichthyes:

Rajidae). Copeia 1998(2):271-290.

g

ISO [International Organization for Standardization]. 1981. ISO standards handbook 3: statistical methods. 2nd ed. Geneva, Switzerland: ISO;

449 p.
Page iii

Contents

Abstract ...............................................................................................................................................................................vii

Introduction ........................................................................................................................................................................... 1

Methods ................................................................................................................................................................................ 1

Results and Discussion ......................................................................................................................................................... 2

Diets ................................................................................................................................................................................ 2

Winter Flounder ....................................................................................................................................................... 2

Windowpane ............................................................................................................................................................ 4

Little Skate ............................................................................................................................................................... 5

Scup ......................................................................................................................................................................... 5

Summer Flounder ..................................................................................................................................................... 6

Red Hake .................................................................................................................................................................. 7

Weakfish .................................................................................................................................................................. 7

Spotted Hake ........................................................................................................................................................... 8

Striped Searobin ....................................................................................................................................................... 8

Northern Searobin .................................................................................................................................................... 9

Striped Bass ........................................................................................................................................................... 10

Clearnose Skate ...................................................................................................................................................... 10

Bluefish .................................................................................................................................................................. 11

Winter Skate ........................................................................................................................................................... 11

Black Sea Bass ....................................................................................................................................................... 12

Spot........................................................................................................................................................................ 12

American Lobster................................................................................................................................................... 13

Tautog .................................................................................................................................................................... 13

Smooth Dogfish ..................................................................................................................................................... 14

Silver Hake ............................................................................................................................................................. 14

Less Abundant Predators ...................................................................................................................................... 15

Overall Perspective on Diets .................................................................................................................................. 16

Forage Base .................................................................................................................................................................. 16

Dominant Invertebrate Prey and Their Life Histories and Habitats ....................................................................... 17

Habitat and Community Associations of Invertebrate Prey .................................................................................. 19

Fish as Prey and Their Life Histories and Habitats ................................................................................................ 20

Overall Perspective on Forage Base ...................................................................................................................... 22

Acknowledgments ............................................................................................................................................................... 22

References Cited .................................................................................................................................................................. 22

Tables

Table 1. Habitat characteristics of strata sampled in the Hudson-Raritan Estuary, 1992-97 .......................................... 28

Table 2a. Summary of Hudson-Raritan Estuary winter flounder diet by sampling period during 1996-97 ...................... 30

Table 2b. Summary of Hudson-Raritan Estuary winter flounder diet by predator size group during 1996-97 ................. 31

Table 2c. Summary of Hudson-Raritan Estuary juvenile winter flounder diet by sampling period during 1996-97 ......... 32

Table 2d. Summary of Hudson-Raritan Estuary adult winter flounder diet by sampling period during 1996-97 ............. 32

Table 2e. Summary of Hudson-Raritan Estuary juvenile winter flounder diet by sampling stratum during 1996-97 ....... 33

Table 2f. Summary of Hudson-Raritan Estuary adult winter flounder diet by sampling stratum during 1996-97 ........... 33

Table 3. Summary of other winter flounder diet studies for the coastal Middle Atlantic Bight .................................... 34

Table 4a. Summary of Hudson-Raritan Estuary windowpane diet by sampling period during 1996-97 .......................... 36

Table 4b. Summary of Hudson-Raritan Estuary windowpane diet by predator size group during 1996-97 ..................... 37

Table 4c. Summary of Hudson-Raritan Estuary windowpane diet by sampling stratum during 1996-97 ........................ 38

Table 5. Summary of other windowpane diet studies for the coastal Middle Atlantic Bight ........................................ 38

Table 6. Summary of Hudson-Raritan Estuary little skate diet by sampling period during 1996-97 .............................. 40

Table 7. Summary of other little skate diet studies for the coastal Middle Atlantic Bight ............................................ 40

Table 8. Summary of Hudson-Raritan Estuary scup diet by sampling period during 1996-97 ....................................... 42
Page iv

Table 9. Summary of other scup diet studies for the coastal Middle Atlantic Bight ..................................................... 42

Table 10. Summary of Hudson-Raritan Estuary summer flounder diet by sampling period during 1996-97 .................... 44

Table 11. Summary of other summer flounder diet studies for the coastal Middle Atlantic Bight .................................. 44

Table 12. Summary of Hudson-Raritan Estuary red hake diet by sampling period during 1996-97 ................................. 46

Table 13. Summary of other red hake diet studies for the coastal Middle Atlantic Bight ............................................... 46

Table 14. Summary of Hudson-Raritan Estuary weakfish diet by sampling period during 1996-97 ................................ 48

Table 15. Summary of other weakfish diet studies for the coastal Middle Atlantic Bight............................................... 48

Table 16. Summary of Hudson-Raritan Estuary spotted hake diet by sampling period during 1996-97 .......................... 50

Table 17. Summary of other spotted hake diet studies for the coastal Middle Atlantic Bight ........................................ 50

Table 18. Summary of Hudson-Raritan Estuary striped searobin diet by sampling period during 1996-97 ..................... 52

Table 19. Summary of other striped searobin diet studies for the coastal Middle Atlantic Bight ................................... 52

Table 20. Summary of Hudson-Raritan Estuary northern searobin diet by sampling period during 1996-97 .................. 54

Table 21. Summary of other northern searobin diet studies for the coastal Middle Atlantic Bight ................................ 54

Table 22. Summary of Hudson-Raritan Estuary striped bass diet by sampling period during 1996-97 ........................... 56

Table 23. Summary of other striped bass diet studies for the coastal Middle Atlantic Bight ......................................... 56

Table 24. Summary of Hudson-Raritan Estuary clearnose skate diet by sampling period during 1996-97 ...................... 58

Table 25. Summary of other clearnose skate diet studies for the coastal Middle Atlantic Bight .................................... 59

Table 26. Summary of Hudson-Raritan Estuary bluefish diet by sampling period during 1996-97 .................................. 60

Table 27. Summary of other bluefish diet studies for the coastal Middle Atlantic Bight ................................................ 60

Table 28. Summary of Hudson-Raritan Estuary winter skate diet by sampling period during 1996-97 ........................... 62

Table 29. Summary of other winter skate diet studies for the coastal Middle Atlantic Bight ......................................... 63

Table 30. Summary of Hudson-Raritan Estuary black sea bass diet by sampling period during 1996-97 ....................... 64

Table 31. Summary of other juvenile black sea bass diet studies for the coastal Middle Atlantic Bight ........................ 64

Table 32. Summary of Hudson-Raritan Estuary spot diet by sampling period during 1996-97 ....................................... 66

Table 33. Summary of other spot diet studies for the coastal Middle Atlantic Bight ..................................................... 66

Table 34. Summary of Hudson-Raritan Estuary American lobster diet by sampling period during 1996-97 ................... 68

Table 35. Summary of other American lobster diet studies for the coastal Middle Atlantic Bight ................................. 69

Table 36. Summary of Hudson-Raritan Estuary tautog diet by sampling period during 1996-97 .................................... 70

Table 37. Summary of other tautog diet studies for the coastal Middle Atlantic Bight .................................................. 70

Table 38. Summary of Hudson-Raritan Estuary smooth dogfish diet by sampling period during 1996-97 ..................... 72

Table 39. Summary of other smooth dogfish diet studies for the coastal Middle Atlantic Bight ................................... 72

Table 40. Summary of Hudson-Raritan Estuary silver hake diet by sampling period during 1996-97 ............................. 74

Table 41. Summary of other juvenile-or-older silver hake diet studies for the coastal Middle Atlantic Bight ................ 74

Table 42. Most commonly eaten invertebrate prey in the Hudson-Raritan Estuary by the 20 most common predators

examined in this survey ................................................................................................................................... 76

Table 43. Habitat associations of other, less commonly eaten invertebrate prey in the Hudson-Raritan Estuary ......... 78

Table 44. Most commonly eaten fish prey in the Hudson-Raritan Estuary .................................................................... 80

Figures

Figure 1. Upper Panel: Habitat features of the Hudson-Raritan Estuary. Lower Panel: Locations of sampling

strata (#s 1-9) and blocks in the Hudson-Raritan Estuary. .............................................................................. 83

Figure 2. Winter flounder distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 ............................. 84

Figure 3. Windowpane distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 ................................. 85

Figure 4. Little skate distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 ..................................... 86

Figure 5. Scup distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 .............................................. 87

Figure 6. Summer flounder distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 ........................... 88

Figure 7. Red hake distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 ....................................... 89

Figure 8. Weakfish distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 ....................................... 90

Figure 9. Spotted hake distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 ................................. 91

Figure 10. Striped searobin distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 ............................ 92

Figure 11. Northern searobin distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 ......................... 93

Figure 12. Striped bass distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 .................................. 94

Figure 13. Clearnose skate distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 ............................. 95

Figure 14. Bluefish distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 ......................................... 96
Page v

Figure 15. Winter skate distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 .................................. 97

Figure 16. Black sea bass distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 ............................... 98

Figure 17. Spot distribution and relative abundance in Hudson-Raritan Estuary, 1992-97............................................... 99

Figure 18. American lobster distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 ......................... 100

Figure 19. Tautog distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 ......................................... 101

Figure 20. Smooth dogfish distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 ........................... 102

Figure 21. Silver hake distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 ................................... 103

Figure 22. Atlantic rock crab distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 ........................ 104

Figure 23. Lady crab distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 .................................... 105

Figure 24. Bay anchovy distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 ............................... 106

Acronyms

DO = dissolved oxygen

ES = empty stomachs

FL = fork length

FO = mean percent frequency of occurrence

IRI = index of relative importance

TDW = mean percent contribution to total stomach content dry weight

TL = total length

TN = mean percent contribution to total number of individual items in the stomach

TV = mean percent contribution to total stomach content volume

TW = mean percent contribution to total stomach content weight

YOY = young of the year

Page vii

ABSTRACT

Characterizing the demersal fish food web in the Hudson-Raritan Estuary is important for understanding specifically

how this estuary is used by fishery resources. Knowledge of fish food webs and essential fish forage resources of the

estuary can support habitat management decisions. Little is known about diets of the community of fish and the American

lobster (Homarus americanus) that inhabit this estuary, although it is a major estuarine complex in the Northeast that

continues to support fisheries. To gain insight into trophic and habitat functions in this estuary, the diets of the most

abundant demersal fish species and the American lobster were examined. These predators were collected by trawl in various

parts of the Hudson-Raritan Estuary over six seasons, July 1996 through November 1997.

The most widely preyed-upon taxa were crustaceans, such as: small or juvenile decapods (e.g., sevenspine bay or

sand shrimp (Crangon septemspinosa), hermit crabs (Pagurus spp.), juvenile Atlantic rock crabs (Cancer irroratus), lady

crabs (Ovalipes ocellatus), and mud crabs (Xanthidae)); the mysid Neomysis americana; and several amphipod species.

Clam siphons, primarily from the northern quahog (Mercenaria mercenaria) and Atlantic surfclam (Spisula solidissima),

were commonly preyed upon by winter flounder (Pseudopleuronectes americanus), as well as by scup (Stenotomus chrysops)

and spot (Leiostomus xanthurus) during some seasons. The diets of common fish and the American lobster in the humanstressed Hudson-Raritan Estuary are similar to those in other, less-stressed estuaries in the Middle Atlantic Bight.

Page 1

INTRODUCTION

The Lower Hudson-Raritan Estuary (hereafter, the Estuary), located at the mouths of the Hudson, Raritan, and

Navesink-Shrewsbury Rivers (New Jersey - New York), is

the polyhaline part of a major, urban, estuarine complex in

the Northeast. The Estuary has supported diverse and productive commercial and recreational fisheries (MacKenzie

1992). Many of these fisheries are gone or operate at a

reduced level because of low resource abundances, harvest regulations, and/or habitat degradation.

The Estuary has been characterized as one of the most

human-altered on the East Coast (Wolfe et al. 1996). Although some sources of habitat alteration or degradation in

the Estuary (e.g., point-source discharges and marsh filling) are being largely controlled through regulation, other

sources (e.g., nonpoint-source discharges and toxic substance spills) continue with little effective control, and new

activities have the potential for adverse effects (Palermo et

al.1998). Despite these alterations, the Estuary is still used

by a diversity of aquatic species (Wilk et al.1998).

To conserve and restore the Estuary’s fishery resources,

there is a need for community- or ecosystem-level information on the status and function of the Estuary’s various

habitats and associated species to provide advice for policy

decisions on conflicting uses of the Estuary. Characterizing fish and American lobster diets in the Estuary is critical

for understanding the value and habitat sources of various

prey taxa in the estuarine food web. Knowledge of food

webs and key predator-prey relationships is important for

habitat-use policy development (Hartman and Brandt 1995).

Although broadscale trophodynamic studies have been

conducted in many other Middle Atlantic Bight estuaries,

e.g., Long Island Sound (Richards 1963), central New Jersey (Festa 1979), Delaware Bay (de Sylva et al.1962), and

Chesapeake Bay (Homer and Boynton 1978), as well as offshore in the New York Bight (e.g., Sedberry 1983; Bowman

et al.1987), the Hudson-Raritan Estuary has never had such

an effort. Only the middle Hudson River part of the Estuary

(near Indian Point, New York) has received attention for

general dietary analysis (Gladden et al.1988), although there

have been focused dietary analyses of a few species such

as striped bass (Morone saxatilus) and juvenile bluefish

(Pomatomus saltatrix). Also, Stehlik et al.(in preparation)

examined the diets of several species of crabs within Raritan

Bay, which complements the present study. Little else has

been reported on the diets of the demersal fishery resource

community of the Estuary, except for some brief incidental

or anecdotal observations (Hall 1894; Merrill 1904; Breder

1922b; NJDEP 1975; Lynch et al.1977; Lawler, Matusky &

Skelly Engineers 1980; Conover et al.1985).

To address this information deficiency, we report on

the results of a seasonal study of the diets of common demersal fish species and the American lobster (Homarus

americanus) collected in various parts of the Estuary. This

study is roughly modeled on Festa’s (1979) study for a shallow, south New Jersey estuary, and is intended to complement that effort, as well as the cursory dietary information

in Able and Fahay (1998). These results are also compared

to a comprehensive summary of most other dietary studies

for the same predators in other Middle Atlantic Bight estuarine or coastal areas. A brief summary of the life history

and habitat of major prey is also included because many of

the habitat issues that managers have to deal with involve

potential perturbations to the health and availability of common prey. This report is intended to be a ready source of

trophic and habitat-use information for subtidal habitat

management within this estuary.

METHODS

The strata and blocks (areas) that were sampled to collect fish and American lobster for stomach content analysis

covered most of the Estuary (Figure 1), but were restricted

to depths greater than 3.0 m because of survey vessel operational factors. The habitat characteristics of these strata

are summarized in Table 1.

Six seasonal sampling periods were used to collect specimens for diet analysis: 1) July 8-12, 1996; 2) October 7-10,

1996; 3) January 27-30, 1997; 4) April 22-29, 1997; 5) August

18-28, 1997; and 6) November 17-20, 1997. In addition, a

special collection of scup (Stenotomus chrysops) was made

during June 9-11, 1997; data on scup from that collection

are included in the August 18-28, 1997, sampling period.

For each sampling period, approximately 40 blocks were

randomly selected from about 200 possible blocks within

the nine sampling strata.

Fish and American lobster samples were collected by a

semiballoon otter trawl that had a 8.5-m headrope, 10.4-m

footrope, 10.2-cm stretch-mesh nylon net, and a 3.5-cm

stretch-mesh liner in the cod end. This trawl was towed for

15 min at ~3.7 km/hr (2 knots). Hydrographic data (i.e.,

depth, salinity, temperature, and dissolved oxygen) were

collected after each successful tow using a “Hydrolab Surveyor 4” multisensor. [Use of trade names is for information

only, and does not represent endorsement by NMFS.] Details of the overall trawl survey are available in Wilk et

al.(1998).

After the trawl was retrieved, the catch was sorted to

species, weighed (g), and measured (0.1 cm). Then, up to

about 10-15 specimens of each nonplanktivorous fish species were selected for analyses, as available. If available,

additional samples were also collected for each apparent

size class of a species. As feasible, the stomachs of large

fish such as skates, dogfishes, and adult striped bass were

examined in the field, or the eviscerated stomachs of such

fish were placed individually in labeled plastic bags and

quickly frozen. Small specimens were also bagged and frozen whole for later laboratory analysis.

To examine the diets in the field or laboratory, the contents of each stomach were carefully emptied onto a gridded
Page 2

petri dish. The total stomach bolus volume was visually

estimated by a side-by-side comparison with a set of variable-diameter, volume-calibrated (cm3

) cylinders. The bolus was separated and examined (by dissecting microscope,

if necessary), then the stomach items or prey were segregated into the lowest identifiable taxon, counted, and measured for length (if possible), and finally, the proportion of

each prey taxon or other item to the total stomach volume

was estimated visually using the petri dish grid. Items or

prey were identified to the lowest level practical using numerous taxonomic references, e.g., Bigelow and Schroeder

(1953), Gosner (1973), and Weiss (1995). The findings of

clam siphons in winter flounder (Pseudopleuronectes

americanus) and a few other predators prompted the collection of whole specimens of larger bivalve mollusks which

commonly occurred in the area in order to examine their

siphons for characteristics that could identify the specific

source of the siphons that were found in the stomachs.

These characteristics were used to develop a rough guide

to siphons to improve the level of prey species identification.

The young-of-the-year (YOY) stages of most fish species (either as predator or prey) examined in the analyses

were identified mostly using Bigelow and Schroeder (1953),

Fitz and Daiber (1963), and a prepublication draft of Able

and Fahay (1998). The transition lengths at 50% maturity,

used to segregate juvenile and adults of certain fish species as part of the diet analysis, were based on O’Brien et

al. (1993).

A literature review of target predator diets in the coastal

Middle Atlantic Bight, the area between Cape Cod and Cape

Hatteras, was used to create summary tables of the diet of

each predator for comparison with results of the present

study. In these tables, prey were listed by their relative

overall importance using several ranking metrics, as available from the document source: mean percent frequency of

occurrence (FO), mean percent contribution to total stomach content volume (TV), mean percent contribution to total stomach content weight (TW), mean percent contribution to total stomach content dry weight (TDW), mean percent contribution to total number of individual items in the

stomach (TN), and an index of relative importance (IRI; FO

x TV, or, FO x TW). Those comparative data available as FO

and TV are the same as those used in the present study.

The two “fresh condition” variables of TV and TW are nearly

equivalent (i.e., 1 g of prey as “fresh” weight approximates

1 cm3

 of prey as “fresh” volume) for most prey such as

crustaceans, polychaetes, fish, shell-less mollusk meat, etc.,

but not for heavy-shelled prey (e.g., sand dollars and mollusks in the shell) consumed whole (Steimle et al.1994).

Our results for diets focus on dominant prey used by

predators found in the Estuary. Dominant prey are generally defined as those contributing five or more percent to

total stomach content volume, but prey of fishery management significance, such as juvenile fishery species, are also

noted. The results are also presented in the order of predator sample abundance. For each predator, the results of this

study are followed by the summary of the results of other

studies for comparative purposes. Because of the relatively large number of predators considered in this predator-community-focused report, this strategy of reporting

by predator sample abundance keeps relevant information

together for each predator, and should be most convenient

to users of this report. Summaries focused on common

prey, with a brief review of their life history and habitat

associations, are also presented.

RESULTS AND DISCUSSION

DIETS

Winter Flounder (Pseudopleuronectes

americanus)

Hudson-Raritan Estuary Results

This species is a common, year-round inhabitant of the

Estuary, and was collected in a range of sizes (6.1-45.0 cm

total length (TL), mean of 20.0 cm) and strata, except in the

central and coastal parts of Lower Bay, Stratum 3 (Figure 2).

The 710 winter flounder which were examined ate 80 distinct, identifiable prey taxa, although only about 20 benthic

invertebrates occurred at a relatively high FO. Endobenthic

and epibenthic polychaetes (21+ species), amphipods (14+

species), and mollusks (10+ species) dominated the diet.

This flounder also ate a range of food types, from plant

detritus to algae to tunicates, including planktonic copepods (e.g., Pseudodiaptomus coronatus), suprabenthic

mysids and the amphipod Gammarus lawrencianus, as well

as the epibenthic and endobenthic invertebrates. Typically, smaller species, earlier life stages, and/or fragments of

larger benthic species were eaten. Some larger bivalve mollusks, e.g., northern quahogs (Mercenaria mercenaria) and

Atlantic surfclams (Spisula solidissima), were important in

the diet, but only their siphons were nipped or torn off by

this nearly toothless predator. The blue mussels (Mytilus

edulis) that were found in the diet, e.g., during April 1997

(Table 2a), were all spat less than 1 cm in length. The decapod crabs that were eaten – Atlantic rock crab (Cancer

irroratus), blue crab (Callinectes sapidus), lady crab

(Ovalipes ocellatus), and Libinia sp. – were all juveniles.

A diversity of tube-dwelling amphipods were also eaten,

especially Ampelisca abdita. Although several polychaete species were identified as being eaten, only the tubedwelling Asabellides oculata and Sabellaria vulgaris, and

the blood worm Glycera sp., were relatively common in the

diet, i.e., occurring in the top 20 prey ranked by FO (Table

2a). The percentage of empty stomachs that were found

ranged from 2.9% in April 1997 to 42.2% in January 1997;

this variable generally ranged between 6.7 and 16.9% for

other sampling periods (Table 2a).
Page 3

Unidentified organic matter (i.e., detritus) ranked as the

most frequently occurring diet item, followed by northern

quahog siphons, Ampelisca abdita, Atlantic surfclam siphons, and unidentified polychaetes or their fragments

(Table 2a). The TV of prey or prey type for all samples was

again dominated by unidentified organic matter, Atlantic

surfclam and northern quahog siphons, the mysid Neomysis

americanus (hereafter, “Neomysis”), unidentified clam siphons, A. abdita, and unidentified polychaetes (Table 2a).

Other prey individually represented a TV of less than 3%.

As with FO, there was a high degree of intersample variability (Table 2a).

For most prey, there were only small seasonal variations in the degree of their use by winter flounder, but for

some prey, there were obvious differences. For example,

there was minimal use of clam siphons during January 1997

(Table 2a). At the same time, there was increased use of

Neomysis and nemerteans. Seasonal predation peaks for

other prey varied annually, i.e., there was relatively high

predation during one summer sampling, but not the other

summer sampling, covered in this survey (e.g., predation

on juvenile Atlantic rock crabs and Asabellides oculata;

see Table 2a).

Other prey or items found in winter flounder stomachs

in lesser quantities were: green and red algae; anthozoans;

nematodes; bryozoans; gastropods (juvenile Crepidula sp.,

Lacuna vincta, Epitonium sp., Astyris lunata, and

Nassarius trivittatus); bivalve mollusks (Solemya velum,

Nucula sp., Mulinia lateralis, Tellina agilis, softshell (Mya

arenaria) siphons, and Lyonsia hyalina); polychaetes

(Phyllodoce sp., Eteone sp., unidentified polynoids, Nephtys

sp., Nereis succinea, N. grayi, Nereis sp., unidentified

capitellids, Asychis elongata, Clymenella torquata,

Spiochaetopterus oculatus, Sabellaria vulgaris, Diopatra

cuprea, Lumbrineris sp., Arabella iricolor, Pectinaria

gouldii, Melinna cristata, Nicolea venustula, and Pherusa

affinis); arachnids (juvenile Limulus polyphemus); copepods (unidentified calanoids, harpacticoids, and cyclopoids,

and the calanoid Pseudodiaptomus coronatus); cumaceans

(Diastylis sp.); tanaids (unidentified); isopods (Edotea

triloba and Cyathura sp.); amphipods (Lembos websteri,

Ericthonius sp., Gammarus sp., Jassa falcata, Hippomedon

serratus, Orchomenella sp., Photis sp., Phoxocephalus

holbolli, Stenothoe sp., and Parametopella sp. (cypris?));

mysids (Heteromysis formosa); decapod crustaceans

(Pagurus sp., P. longicarpus, xanthids (Dyspanopeus?),

juvenile blue crabs, juvenile Libinia sp., and juvenile

Ovalipes ocellatus); echinoderms (juvenile Echinarachnius

parma); tunicates (Molgula sp.); and sand, shell hash, organic detritus, and manmade artifacts such as coal granules

and synthetic fibers.

Winter flounder diet changed with size/growth. This

shift in use of common prey was generally from small crustaceans (mysids and amphipods), polychaetes, and detritus by smaller fish, to more bivalve mollusk siphons by

larger fish (Table 2b).

Winter flounder diet was examined for seasonal shifts

in prey use as related to flounder size and maturity. Because of small sample sizes for each of the four size groups

portrayed in Table 2b, the samples were pooled into two

groups: juvenile (less than 20 cm TL; Table 2c) and adult

(greater than or equal to 20 cm TL; Table 2d). In the summer-fall, juvenile winter flounder focused their feeding on

northern quahog and Atlantic surfclam siphons, an amphipod (i.e., Ampelisca abdita or A. vadorum), a tube-dwelling

polychaete (Sabellaria vulgaris), and detritus, although

Neomysis became important as prey in the winter (Table 2c).

Other prey were relatively evenly used during most seasons or showed no seasonal pattern of use. For adults, the

list of commonly eaten prey was condensed, with only four

distinct prey being notable, and seasonal sample sizes were

more irregular and often inadequate (Table 2d). Again, clam

siphons were the dominant prey. The large bloodworm

Glycera sp. and juvenile Atlantic rock crabs were the only

other prey with any seasonal peaks.

Some studies of the winter flounder diet have shown

that the diet closely reflects environmental conditions and

prey availability in the areas in which the fish are collected

(Frame 1974; MacPhee 1969). The winter flounder diet in

this study also showed differences in prey use that varied

among sampling strata, although sample sizes were small

for some strata, particularly for the channel habitats, Strata

7-9 (Tables 2e,f). Some prey (e.g., Glycera sp.) were eaten in

similar proportion by juveniles and adults from the same

strata areas of the Estuary. Other prey (e.g., Neomysis,

Crangon septemspinosa (hereafter, “Crangon”), and

Gammarus sp.) were found in stomachs of juveniles or

adults, but not both. The prey of juvenile winter flounder

among different strata suggest no habitat-related patterns

(Table 2e). There was no specific association of prey with

channels (Strata 7-9), nor with the western or eastern areas,

with the possible exception of the polychaetes Glycera sp.

and Sabellaria vulgaris and northern quahog siphons in

the western Strata 1-3, and Atlantic surfclam siphons in

eastern Strata 4 and 6 (Table 2e). Predation by juveniles

and adults on northern quahog siphons appears restricted

to less-saline, western Strata 1-3 and 6, and predation on

Atlantic surfclam siphons occurs in marine eastern Strata 4

and 5 and in channel Strata 7 and 8, as might be expected

from the Atlantic surfclam’s salinity preferences.

Comparisons with Other Diet Studies

Other winter flounder dietary studies in and near the

Estuary, or within the coastal Middle Atlantic Bight, found

a similar diet to that reported here, i.e., opportunistic predation on benthic invertebrate macrofauna, especially polychaetes and amphipods, and on zooplankton by the smallest winter flounder sizes (Table 3). However, there was an

unusually high degree of molluscan siphon nipping in this

estuary compared to what has been reported elsewhere
Page 4

(Tables 2 & 3; Lawler, Matusky & Skelly Engineers 1980;

Stehlik and Meise 2000). A lesser degree of molluscan siphon nipping by winter flounder has been also reported in

Canada (Medcoff and McPhail 1952), Cape Cod Bay (Gilbert and Suchow 1977), Long Island Sound (Carlson 1991),

and near Woods Hole, Massachusetts (Frame 1974; Lux et

al.1996), and in a few other studies (Table 3), however. It

has been assumed that this siphon nipping is nonlethal to

the mollusks, which can be fishery resources in their own

right, and that the siphons regenerate (Irlandi and Mehlich

1996).

Windowpane (Scophthalmus aquosus)

Hudson-Raritan Estuary Results

Windowpane from YOY to adult (range of 2.5-35.0 cm

TL, mean of 20.7 cm) are a year-round inhabitant of this

estuary. Five hundred seventy windowpane were examined from all areas, although slightly greater quantities were

available from the channels, especially in and near Raritan

Channel, Stratum 9 (Figure 3). At least 37 prey taxa were

identified in their diet. This prey spectrum included two

mysid species, three or more decapod crustacean species,

seven amphipod species, two or more copepod species,

eight mollusk species, nine identifiable species of larval or

juvenile fish, and some miscellaneous, nonprey items (green

algae to coal fragments). This prey spectrum included a mix

of benthic, suprabenthic, and pelagic species.

Despite the overall diversity of prey consumed, windowpane have a relatively focused diet. By FO for all

samples, Neomysis was the dominant prey at 65.9% (range

of 33.7-93.3%). It was followed in importance by Crangon

at 31.7% (range of 23.6-53.0%) and the suprabenthic amphipod Gammarus lawrencianus at 9.5% (range of 0.8-39.0%).

The other prey were eaten at a low FO (i.e., less than 5%).

The percentage of empty stomachs ranged from 2.0% in

July 1996 to 33.7% in January 1997, and was between 10 and

24% in other sampling periods (Table 4a).

The TV paralleled the FO. Neomysis made up 57.1% of

the overall diet by TV (range of 17.8-70.1%), with Crangon

contributing 29% (range of 21.3-47.7%). Most other prey

individually represented less than 0.1% of TV, although

higher values occurred during some sampling periods (Table

4a).

Other prey or items found in windowpane stomachs in

lesser quantities were: green algae; hydroids; nemerteans;

gastropods (Lacuna vincta, juvenile Crepidula sp.,

Nassarius trivittatus, and Astyris lunata); bivalve mollusks

(Mulinia lateralis, Nucula sp., unidentified, and blue mussel spat); cephalopods (unidentified squid); polychaetes

(unidentified); copepods (unidentified); cumaceans (unidentified); amphipods (Corophium sp., Jassa falcata,

Stenothoe sp., Hippomedon serratus, and Unciola sp.);

mysids (Heteromysis formosa); decapod crustaceans

(Pagurus longicarpus, Palaemonetes vulgaris, and unidentified zoea); fish (unidentified juvenile flounder, unidentified juvenile fish, juvenile Atlantic menhaden (Brevoortia

tyrannus), juvenile herring (Alosa), juvenile red hake

(Urophycis chuss), juvenile cunner (Tautogolabrus

adspersus), Menidia sp., juvenile Atlantic croaker

(Micropogonias undulatus), and juvenile sand lance

(Ammodytes sp.)); and sand and coal pebbles.

There was seasonal variability in the consumption of

some prey (e.g., the use of Neomysis peaked during the

summer). The use of Crangon peaked in the winter-spring,

although it was a major prey in July 1996 samples (Table 4a).

G. lawrencianus was mostly consumed in the fall, especially in 1996, as was the red copepod Pseudodiaptomus

coronatus.

There was a clear shift evident in prey use with windowpane growth (e.g., from Neomysis as overwhelmingly

dominant for windowpane less than 20.0 cm TL, to Crangon

for windowpane at larger sizes, although Neomysis was

dominant at all sizes (Table 4b). Small fish (e.g., Anchoa sp.)

also become more important for the larger-sized fish.

The results suggest some differences in prey use among

strata and regions (Table 4c). Neomysis seems to be the

basic prey in the Ambrose Channel to Verrazano Narrows

area (Strata 6 and 7). Crangon, on the other hand, seems to

be more often eaten in the central-western areas of the Estuary (i.e., Strata 2, 3, 8, and 9). Other prey constituted an

insignificant proportion (a TV of less than 10%) of the diet

in most areas, except in Strata 4 and 5 (the marine shoals)

where small or juvenile fish were eaten. Windowpane abundances were highest in channels (Figure 3) where Neomysis

might be most abundant; see the “Forage Base” section for

a discussion of the habitat of Neomysis.

Comparisons with Other Diet Studies

No previous focused studies of the diet of this species

are known for this estuary, although Breder (1922b) commented that stomach contents of a few specimens that he

examined “consisted of crustacean remains, probably

schizopods [mysids].” In general, other studies of the diet

of this species in the Middle Atlantic Bight found mysids

(especially Neomysis), Crangon, and “nekton” (i.e., small

fish and squid) to be primary prey (Table 5), but smallersized windowpane also ate copepods. For the continental

shelf, Langton and Bowman (1981) reported 40-60% of the

windowpane that they had examined had empty stomachs,

but mysids and shrimp continued to dominate the diet offshore. The results of the present study are consistent with

those of other studies and with Bigelow and Schroeder’s

(1953) general summary of the diet, except for the relatively

high use of G. lawrencianus as prey in the present study.
Page 5

Little Skate (Raja erinacea)

Hudson-Raritan Estuary Results

Little skate (mostly adults) were commonly found

throughout this estuary during most seasons, except the

summer (Figure 4; Wilk et al.1998). The stomachs of 332

little skate (range of 33.0-49.0 cm TL, mean of 43.2 cm) were

examined. Over 50 prey taxa or items, which ranged from

green algae to a variety of small fish, were identified in the

little skate diet. These prey taxa included 11 decapod crustaceans, 6 amphipods, 5 polychaetes, 8 mollusks, 10 identifiable fish, and miscellaneous prey or items (Table 6).

The most frequently found prey, overall, was Crangon

at an FO of 82.8% (range of 77.2-92.9%). This prey was

followed by juvenile or small Atlantic rock crabs at an FO of

49.5% (range of 7.1-75.3%), which were often found in a

soft-shell stage, then by Neomysis at an FO of 16.3% (range

of 6.1-28.5%) and Ovalipes ocellatus at an FO of 10.9%

(range of 1.2-36.4%). The remaining prey had overall FOs

of less than 10% (Table 6), with few empty stomachs.

The TV parallels the FO, with Crangon having 29.6%

(range of 20.3-90.1%) of the TV, Atlantic rock crabs having

18.6% (range of 4.7-38.1%), and other prey having less than

10%, with the exception of the October 1996 sampling period when O. ocellatus had 15.6% (Table 6). A number of

juvenile blue crabs were also eaten (Table 6).

Other prey or items found in little skate stomachs in

lesser quantities were: unidentified green algae; hydroids;

nematodes; nemerteans; gastropods (Lacuna vincta,

Nassarius trivittatus, and Astyris lunata); bivalve mollusks

(blue mussels, Mulinia lateralis, northern quahog siphons,

and unidentified); polychaetes (Lumbrineris sp.,

Spiochaetopterus oculatus, Arabella iricolor, Pherusa

affinis, Diopatra cuprea, and unidentified); copepods

(Pseudodiaptomus coronatus); cumaceans (unidentified);

isopods (Cirolana concharum, and Cyathura sp.); amphipods (Leptocheirus pinguis, Hippomedon serratus, Unciola

sp., Ampelisca abdita, and unidentified); decapod crustaceans (unidentified, xanthids, Pagurus pollicaris, P.

longicarpus, Palaemonetes vulgaris, Dichelopandalus

leptocerus, and Axius serratus); stomatopods (Squilla

empusa); fish (Raja sp. egg case fragments, juvenile rock

gunnel (Pholis gunnellus), juvenile windowpane, northern

searobin (Prionotus carolinus), unidentified searobins,

sand lance, smallmouth flounder (Etropus microstomus),

goby (Gobiosoma sp.), northern pipefish (Syngnathus

fuscus), juvenile red hake, juvenile winter flounder, and juvenile silver hake (Merluccius bilinearis)); and sand, wood

fragments, and human artifacts such as coal granules, iron

rust flakes, and plastic particles.

Although little skate tended to be most common in this

estuary in the cooler months, and sample sizes are relatively small in the summer, there appears to be a possible

predation emphasis on Atlantic rock crabs and Neomysis

during the cooler months (Table 6).

Comparisons with Other Diet Studies

No previous studies of the diet of this species are known

for this estuary. However, the diet of this species has been

studied elsewhere, both on the Middle Atlantic Bight continental shelf, and within other bays and estuaries (Table 7).

These studies also show that small crustaceans dominate

the little skate diet; with skate less than 20.0 cm TL eating

small crustaceans (e.g., copepods, mysids, and amphipods

such as Unciola irrorata, Gammarus annulatus,

Leptocheirus pinguis, and Monoculodes edwardsi), and

with larger skate eating more decapod crustaceans, especially Crangon, Cancer sp., Dichelopandalus leptocerus,

and hermit crabs (Pagurus sp.). However, squid and small

fish (e.g., sand lance, butterfish (Peprilus triacanthus), “herring” (Alosa sp.?), searobins, juvenile flounder, and red hake)

were also eaten (Table 7). The diet of little skate from the

Estuary (Table 6) is consistent with these other study results, and with the general dietary summaries reported in

Nichols and Breder (1927) and Bigelow and Schroeder (1953).

Scup (Stenotomus chrysops)

Hudson-Raritan Estuary Results

Scup, mostly juveniles, were found from spring through

fall in the Estuary and were relatively widespread in distribution, although with a tendency to be collected more often in the northern areas (Figure 5). The stomachs of 254

scup (range of 8.0-24.0 cm FL, mean of 12.9 cm) were examined. At least 39 items or prey taxa were identified in their

stomachs, including 8 polychaetes, 7 amphipods, 6 decapod crustaceans, 6 mollusks, 2 mysids, and other taxa (e.g.,

hydroids). The majority of these prey were benthic, except

for mysids and Gammarus lawrencianus (Table 8).

The dominant items in the diet by FO were unidentified

organic matter at 35.8% (range of 21.3-46.1%), Neomysis at

32.3% (range of 17.3-50.0%), bivalve mollusk remains at

14.3% (range of 1.3-26.7%), G. lawrencianus at 16.8% (range

of 0.0-48.3%), Crangon at 15.2% (range of 3.9-30.7%), unidentified polychaetes at 14.2% (range of 6.7-35.5%), and

Ampelisca abdita at 10.1% (range of 0.0-18.0%) (Table 8).

The contribution of prey to the overall TV parallels that

to the overall FO in the same order of contribution (Table 8).

There were few empty stomachs.

Other prey or items found in scup stomachs in lesser

quantities were: unidentified hydroids; unidentified nemerteans; gastropods (juvenile Crepidula sp., unidentified,

and eggs); bivalve mollusks (Tellina agilis and Nucula

sp.); polychaetes (Paranaites speciosa, Asabellides oculata,

Sabellaria vulgaris, Pectinaria gouldii, Phyllodoce sp.,

Pherusa affinis, and Nereis sp.); copepods

(Pseudodiaptomus coronatus and unidentified calanoid);

cirripeds (Balanus sp.); tanaids (unidentified); isopods

(Cyathura sp.); amphipods (unidentified, Orchomenella sp.,
Page 6

Photis sp., caprellids, Unciola sp., and Corophium sp.);

mysids (Mysidopsis bigelowi); decapod crustaceans

(xanthid crabs, juvenile blue crabs, and unidentified); and

fish (silversides (Menidia sp.) and unidentified).

There were few, notable, interannual or seasonal differences in the diet of this basically warm-season species,

with the possible exception of predation on G. lawrencianus

in 1996 that was not evident in 1997 (Table 8).

Comparisons with Other Diet Studies

Within this estuary, the only previous known data on

the scup diet is from the unpublished, preliminary 1976 results of the senior author, who examined 13 juvenile fish

from Strata 1, 3, and 4. He found that the mostly frequently

consumed prey were: the polychaete Asabellides oculata

and copepods in Sandy Hook Bay, polydorid polychaetes

and the dwarf surfclam Mulinia lateralis off Staten Island,

and copepods and blue mussel spat on Romer Shoal (Stratum 4).

Michelman (1988) found that the juvenile scup diet in

Rhode Island varied seasonally, but was still generally focused on benthic invertebrates, such as polychaetes (e.g.,

maldanids, Nephtys sp., Nereis sp., and Pherusa affinis),

small decapod crustaceans (Pagurus sp. and other crabs),

Neomysis, amphipods (Leptocheirus pinguis and others),

as well as mollusks, a burrowing anemone (Ceriantheopsis

americanus), and fish eggs and larvae.

Most other studies found that scup less than 15 cm FL

ate small invertebrates such as copepods, polychaetes,

amphipods, decapod crustaceans (especially juvenile Atlantic rock crabs), and squid (Table 9); a number of qualitative or summary reports have found the same (Baird 1873;

Peck 1896; Nichols and Breder 1927; Hildebrand and

Schroeder 1928; Bigelow and Schroeder 1953; Allen et

al.1978). Linton (1901) and Sedberry (1983) found that the

diet of scup gradually shifted with growth or size from small

pelagic crustaceans to a variety of benthic taxa. The results

of the present study are basically consistent with these

other results, and show a strong reliance on benthic

macrofauna and detritus as prey.

Summer Flounder (Paralichthys dentatus)

Hudson-Raritan Estuary Results

This species was most commonly collected during the

summer and throughout the Estuary, but especially along

the New Jersey shore (Figure 6; Wilk et al.1998). The stomachs of 229 summer flounder (range of 13.8-69.0 cm TL,

mean of 36.0 cm) were examined. Over 35 prey species or

items were identified in their diet, including juvenile or small

adults of 12 species of fish, 5 species of decapod crustaceans, Neomysis, and other taxa (Table 10).

Crangon with an FO of 49.5% (range of 34.4-78.0%)

and Neomysis with an FO of 19.8% (range of 0.0-33.6%)

were most frequently eaten. Unidentified fish were next

with an FO of 13.2% (range of 0.0-14.0%), and juvenile

Ovalipes ocellatus were prominent in the August 1997 stomach samples (Table 10).

The FO ranking was also followed by the TV ranking,

with Crangon having a TV of 29.4%, and Neomysis having

a TV of 11.4% (Table 10). The percentage of empty stomachs ranged from 10 to 50%, with the highest levels being

found in the winter-spring period.

Other prey or items found in summer flounder stomachs in lesser quantities were: unidentified algae; hydroids;

bryozoans; gastropods (Crepidula sp. and Nassarius

trivittatus); bivalve mollusks (blue mussel spat, Mulinia

lateralis, Ensis directus, Nucula sp., and Tellina agilis);

polychaetes (Sabellaria vulgaris and unidentified); copepods (unidentified calanoid); isopods (Cyathura sp.); amphipods (caprellids, Gammarus lawrencianus, and Ampelisca

abdita); decapod crustaceans (juvenile blue crabs, Pagurus

longicarpus, and unidentified); and fish (juvenile scup,

cunner, rock gunnel, juvenile searobins, juvenile weakfish

(Cynoscion regalis), Menidia sp., juvenile striped searobin

(Prionotus evolans), juvenile black sea bass (Centropristis

striata), northern pipefish, juvenile Alosa herring, and juvenile grubby (Myoxocephalus aenaeus)).

Summer flounder were mostly collected in the spring

and summer (Table 10), so seasonal shift could not be examined. There were few notable dietary differences between

1996 and 1997, although in 1997 summer flounder made

greater use of O. ocellatus and unidentifiable juvenile flounder as prey. Most (85%) of the summer flounder examined

were 30 cm TL or more, and probably not YOY. Despite the

small sample size for YOY summer flounder, their diet differed little from that of larger fish: Crangon and Neomysis

dominated the diet, with small Ovalipes and fish being of

notable importance (Table 10).

With the number of samples being few (i.e., less than 30

samples per stratum), and with the distribution of samples

among strata being small, the results of interstrata comparisons were inconclusive, although there was a suggestion

that Crangon were eaten commonly everywhere except in

the Romer Shoal and Ambrose Channel areas (Strata 4 and

7). Neomysis were mostly found in stomachs examined from

deeper channels (Strata 8 and 9) and near the Verrazano

Narrows (Stratum 6), but also within Sandy Hook Bay (Stratum 1). The latter observations perhaps reflected some

recent feeding in the unsampled Sandy Hook and Earl Naval Channels, located between Strata 1, 2, and 4.

Comparisons with Other Diet Studies

No previous studies of the diet of this species are known

for this estuary.
Page 7

In general, YOY summer flounder prey upon small fish

(e.g., silversides, mummichog (Fundulus heteroclitus), bay

anchovy (Anchoa mitchilli), and sticklebacks), and

Palaemonetes, Crangon, and Neomysis shrimps (Table 11).

The species is highly opportunistic, but its diet shifts ontogenetically, from small crustaceans at smaller sizes, to fish

prey at larger sizes. The diet of the predominantly YOY and

juvenile summer flounders examined in the Estuary, dominated by crustaceans and small fish (Table 10), is consistent with other studies (Table 11), and with the generalizations of Hildebrand and Schroeder (1928), Ginsberg (1952),

and Bigelow and Schroeder (1953), which were often based

on small or ambiguous sample sizes.

Red Hake (Urophycis chuss)

Hudson-Raritan Estuary Results

Red hake were commonly collected in channels and the

deep area below the Verrazano Narrows (Gravesend Bay,

Stratum 6; Figure 7) and during most seasons (Wilk et al.1998).

The diet of 166 red hake (range of 4.3-39.0 cm TL, mean of

19.0 cm) was examined. These fish were primarily juveniles

and were most frequently collected during cooler seasons.

At least 33 prey species were identified in the diet, including 7 decapod crustaceans, 9 amphipods, Neomysis, 7 juvenile fishes, and other taxa from algae to mollusks. Most

prey were benthic species.

Crangon with an FO of 77.6% (range of 56.3-100.0%),

Neomysis with an FO of 31.7% (range of 0.0-48.4%),

Gammarus lawrencianus with an FO of 20.9% (range of 0.0-

100.0%), and unidentified organic detritus with an FO of

10.6% (range of 0.0-20.6%) dominated the diet.

Crangon dominated the diet’s TV at 39.0% (range of

23.3-50.0%), followed again by Neomysis at 15.7% (range of

0.0-30.3%) (Table 12). The other prey were infrequently

found in the stomachs, and few stomachs were found empty.

The inadequate samples in 1996 and during warmer months

(Table 12) prevent analysis of seasonal or interannual variation in the diet of this species.

Other prey or items found in red hake stomachs in lesser

quantities were: green algae; hydroids; bivalve mollusks

(Nucula sp., Tellina agilis, blue mussel spat, and unidentified); polychaetes (unidentified and Pherusa affinis); copepods (Pseudodiaptomus coronatus and unidentified

calanoid); isopods (Edotea triloba); amphipods

(Phoxocephalus holbolli, Unciola sp., Ampelisca abdita,

Corophium sp., Jassa falcata, Stenothoe sp., Hippomedon

serratus, and unidentified); decapod crustaceans (juvenile

Libinia sp., Pagurus longicarpus, juvenile Ovalipes

ocellatus, Palaemonetes vulgaris, and unidentified); fish

(juvenile silver hake, juvenile red hake, smallmouth flounder, juvenile searobin, juvenile weakfish, juvenile cunner,

unidentified juvenile flounder, and skate (Raja sp.) egg case

fragments); and wood fragments.

Comparisons with Other Diet Studies

The diet of red hake from the Estuary (Tables 12) is

consistent with other dietary studies for the species, with

crustaceans being primary prey. The only previous, quantitative study of the diet of this species in Raritan Bay examined 45 subadults of this species in spring 1976 within Sandy

Hook Bay and off Staten Island (Steimle, unpubl. data).

That diet was dominated (i.e., a TV of 92-100%) by Crangon.

This result is consistent with Breder’s (1922b) earlier comment that the few red hake that he looked at in Sandy Hook

Bay in summer 1921 were “crammed full of large prawns”;

these “prawns” were further defined as being Crangon in

Nichols and Breder (1927).

In the nearby New York Bight apex (outside the mouth

of the Estuary), over 1,000 red hake were examined and found

to prey most commonly on Crangon, various polychaetes

(mostly Pherusa affinis and Nephtys incisa), Neomysis, and

benthic amphipods (Steimle 1985, 1994) (Table 13).

Hildebrand and Schroeder (1928) observed that red hake

that were caught off Sandy Hook had gorged on sand lance.

In general, the summary of other studies (Table 13) and the

treatise by Bigelow and Schroeder (1953) show that juvenile red hake eat a variety of small benthic and zooplanktonic invertebrates, but primarily crustaceans. Steiner et

al.(1982) reported that juvenile red hake use shelter during

the day (such as living sea scallops, Placopecten

magellanicus) and leave this shelter to feed at night. As

red hake grow, larger crustaceans such as decapods increase in importance in their diet, and some fish are also

eaten (Table 13). Adult red hake were rarely collected within

the Estuary (Wilk et al.1998).

Weakfish (Cynoscion regalis)

Hudson-Raritan Estuary Results

Weakfish were another summer-fall inhabitant of the

Estuary and were collected mostly in or near channels, especially in Stratum 9, Raritan Channel (Figure 8). The stomachs of 197 weakfish (range of 7.5-54.0 cm TL, mean of 17.7

cm) were examined. Over 20 prey species or items were

identified in the diet, but they were dominated by crustaceans and a few juvenile or small fish. Crangon and

Neomysis were the most frequently eaten prey, with only

Gammarus lawrencianus and digested fish (probably bay

anchovy) being of any relative importance (Table 14). There

do not appear to be any consistent interannual differences

in the diet, although there were pulses of the consumption

of Neomysis, bay anchovy, and juvenile silver hake in the

diet during certain sampling periods (Table 14).

Other prey or items found in weakfish stomachs in lesser

quantities were: unidentified hydroids; gastropods

(Nassarius trivittatus); polychaetes (unidentified); crustaceans (unidentified); copepods (Pseudodiaptomus
Page 8

coronatus); amphipods (Corophium sp. and Unciola sp.);

decapod crustaceans (Dichelopandalus leptocerus, juvenile Ovalipes ocellatus, juvenile Atlantic rock crabs, and

juvenile blue crabs); fish (juvenile weakfish, juvenile Atlantic menhaden, butterfish, juvenile unidentified flounder, and

juvenile windowpane); and human artifacts such as cellophane.

Comparisons with Other Diet Studies

The only previous information on the diet of this species known for this estuary are comments by Breder (1922b)

that, when he examined a few adult weakfish from Sandy

Hook Bay, he found that they had eaten fish such as Atlantic menhaden [juveniles?] , silver perch (Bairdiella

chyrsoura), and anchovies, and squid and “prawns”

[Crangon]. Another cursory diet examination by Lynch et

al.(1977) of 25 juvenile fish from the Raritan River (western

boundary of the Estuary) also noted that weakfish there

also ate Crangon and fish. Within the upper Hudson River

Estuary (above Manhattan Island), Gladden et al.(1988) reported that weakfish generally ate “fish and

macroinvertebrates.”

The summaries of the results of other weakfish studies

(Table 15) and the generalized summary of Bigelow and

Schroeder (1953) show that the diet of this species can vary

substantially among estuaries. That is, it can be dominated

by Crangon or small fish (especially bay anchovy and juvenile weakfish) in some estuaries, but by mysids (mostly

Neomysis) or amphipods (e.g., Gammarus sp.) in others.

The earliest studies listed in Table 15 were less precise in

defining prey, but the “shrimp,” “prawns,” or “mysids” that

they noted are almost certainly Crangon and Neomysis,

and suggest that there does not appear to be any substantial shift in dominant prey over the decades, at least in the

past century. Other weakfish diet studies were not listed in

Table 15 because of limitations or the general nature of their

information, e.g., Eigenmann (1902), Linton (1901), Tracy

(1910), Nichols and Breder (1927), Hildebrand and Schroeder

(1928), Lascara (1981), and Grecay (1990). The pattern of

weakfish predation within the Estuary seems to be typical

and focused on both Crangon and Neomysis, but small fish

(e.g., bay anchovy, butterfish, and weakfish) and Gammarus

sp. amphipods are also important (Tables 14 and 15).

Spotted Hake (Urophycis regia)

Hudson-Raritan Estuary Results

Spotted hake of all size and age classes were collected

commonly during the warmer months within the Estuary

and mainly in channels, especially Raritan Channel (Stratum 9, Figure 9). The 162 spotted hake (range of 6.5-33.0 cm

TL, mean of 18.3 cm) which were examined ate 30 prey taxa,

ranging from hydroids to fish. The most frequently eaten

prey were crustaceans (i.e., Crangon, Neomysis, and

Gammarus lawrencianus) and small fish (Table 16). The

copepod Pseudodiaptomus coronatus was frequently eaten

in half of the sampling periods. Few empty stomachs were

found. Crangon dominated the overall stomach volumes

with a TV of 45.7% (range of 31.3-60.9%) (Table 16).

Other prey or items found in spotted hake stomachs in

lesser quantities were: unidentified nematodes; bivalve

mollusks (unidentified, Nucula sp., blue mussel spat, and

juvenile Pitar morrhuanus); polychaetes (unidentified,

Nereis sp., and Pherusa affinis); sipunculids (unidentified);

crustaceans (unidentified); copepods (unidentified); amphipods (unidentified, Ampelisca abdita, Jassa falcata,

Hippomedon serratus, Unciola sp., and Ericthonius sp.);

decapod crustaceans (juvenile Atlantic rock crabs, Ovalipes

oculata, Pagurus sp., Dichelopandalus leptocerus, and

Palaemonetes sp.); and fish (juvenile silver hake, juvenile

red hake, juvenile searobins, and smallmouth flounder).

Comparisons with Other Diet Studies

No previous studies of the diet of this species are known

for this estuary.

In general, other studies show that spotted hake usually eat larger epibenthic crustaceans and small fish (Table

17). Among the crustaceans eaten, Crangon, Neomysis,

copepods, other decapod shrimp, and crabs were prominent in the diet of this species. The variety of fish identified

in these other studies included bay anchovy and sand lance

among others. This diet spectrum is consistent with Bigelow

and Schroeder’s (1953) review, the species’ diet south of

Cape Hatteras (Burr and Schwartz 1986), and with the results of the present study (Table 16).

Striped Searobin (Prionotus evolans)

Hudson-Raritan Estuary Results

Adults and juveniles of this species were collected

mostly during the summer months, and in or near channels

or within Sandy Hook Bay (Figure 10; Wilk et al.1998). The

153 samples of striped searobin (range of 4.5-47.2 cm TL,

mean of 21.4 cm) which were examined ate 34 identifiable

prey taxa, but most frequently preyed upon Crangon,

Neomysis, and other crustaceans, and upon small or juvenile fish. Of interest was the relatively frequent occurrence

of small, approximately 1-3 mm, coal granules or pebbles in
Page 9

their stomachs (Table 18). The TV was dominated (45.8%,

range of 37.4-71.4%) by Crangon (Table 18). The diet was

similar for 1996 and 1997 (Table 18).

Other prey or items found in striped searobin stomachs

in lesser quantities were: unidentified hydroids; gastropods (unidentified, Nassarius obsoletus, and N. trivittatus);

bivalve mollusks (Nucula sp., Mulinia lateralis, and Ensis

directus); cephalopod (unidentified squid); polychaetes (unidentified); crustaceans (unidentified); copepods (unidentified and Pseudodiaptomus coronatus); isopods (Edotea

triloba); amphipods (unidentified, Corophium sp., and

Unciola sp.); mysids (Heteromysis formosa); decapod crustaceans (xanthid crabs and unidentified crab fragments);

fish (smallmouth flounder, juvenile windowpane, juvenile

anchovy, juvenile grubby, unidentified juvenile flounder,

juvenile searobin, juvenile black sea bass, and juvenile stargazer (Astroscopus sp.)); and sand.

Comparisons with Other Diet Studies

There is only one other study of the diet of this species

known for this estuary. Manderson et al. (1999) examined

35 stomachs of this species from shallow water in Sandy

Hook Bay (near Stratum 1) and its Navesink River tributary,

at its southern border. They reported an FO of 68% of YOY

winter flounder in the searobin’s diet, although Crangon

and other crustaceans were the primary prey.

A summary of most other quantitative studies of the

diet of this species from different areas shows that the diet

was also based on crustaceans (e.g., Crangon, Neomysis,

copepods, amphipods, and juvenile crabs) and small or juvenile fish (e.g., winter flounder, striped and northern

searobins, scup, windowpane, bay anchovy, Menidia,

northern pipefish, and probably others) as available (Table

19). Other, more generalized discussions of their diet, e.g.,

Bigelow and Schroeder (1953), also note a broad spectrum

of prey in the diet of this species, including crabs, amphipods, squid, bivalve mollusks, polychaetes, small fish (herring and winter flounder), and algae. In Richards et al.’s

(1979) Long Island Sound study, they reported that the prey

of age 1+ searobin varied with habitat type (i.e., prey eaten

on sandy bottoms were different from prey eaten on muddy

bottoms). For example, on sandy bottoms, the razor clam

(Ensis directus) was important. They also found that some

predation showed no habitat-related differences (e.g., on

Neomysis, Crangon, and Ovalipes ocellatus and other

crabs), and concluded that the diet of the adult striped

searobin when at smaller sizes – although having a great

deal of overlap with the sympatric but smaller northern

searobin – tended to reduce competition for food by focusing on larger prey that were less specific in their habitat

preferences. The results of the present study are consistent with the findings of other studies; although, again, one

or more 2-5 mm diameter pebbles of coal or charcoal were

observed in about 5% (range of 3-17%) of the stomachs

(Table 18).

Northern Searobin (Prionotus carolinus)

Hudson-Raritan Estuary Results

This small species (range of 5.1-20.4 cm TL, mean of

15.1 cm) was collected mostly during the summer, and mainly

in the eastern areas of the Estuary, e.g., between Verrazano

Narrows and Sandy Hook Bay (Figure 11; Wilk et al.1998).

One hundred three northern searobin stomachs were examined, and over 20 prey taxa were identified, which were mostly

crustaceans. The reoccurring prey group of Crangon,

Neomysis, and Gammarus lawrencianus were most frequently found in the stomachs. The contribution of prey to

TV parallels their contribution to FO, although juvenile Atlantic rock crabs were of added importance to the diet of the

large, August 1997 collection sample (Table 20). Coal

pebbles were also found in these stomachs, but only during one collection, and then at an FO of 25% for the 87 fish

examined (Table 20).

Other prey or items found in northern searobin stomachs in lesser quantities were: hydroids (unidentified); nematodes (unidentified); bivalve mollusks (blue mussel spat);

polychaetes (unidentified); copepods (Pseudodiaptomus

coronatus); isopods (Edotea triloba); amphipods

(Corophium sp., Ampelisca abdita, and Leptocheirus

pinguis); mysids (Heteromysis formosa); decapod crustaceans (Pagurus longicarpus); fish (juvenile smallmouth

flounder, juvenile striped searobin, and juvenile black sea

bass); and unidentified organic matter.

Comparisons with Other Diet Studies

No previous studies of the diet of this species are known

for this estuary.

Some results of other studies of the diet of this predator show that, like its sibling species, the striped searobin,

the northern searobin also preys principally upon crustaceans, with Crangon, Neomysis, amphipods, and copepods being prominent in the diet, but fish are eaten to a

lesser degree (Table 21). This dietary pattern was also reported by Hildebrand and Schroeder (1928) and Bigelow

and Schroeder (1953). The smaller adult size of the northern

searobin, compared to the striped searobin, is a logical explanation for the difference in the use of fish (although

juvenile herring, winter flounder, weakfish, bay anchovy,

and others are reported as prey), and perhaps for the slightly
Page 10

greater use of smaller macrofauna such as polychaetes and

cumaceans. In Long Island Sound, Richards et al. (1979)

reported on the diet of YOY and older northern searobins,

and despite some ambiguity in their results, the YOY of this

species appeared to prey principally upon Neomysis and

copepods, based on numbers eaten. Larger fish were more

focused on amphipods, isopods, and small decapod crustaceans as prey. Mann (1974) found that diet of this species varied with sediments and water depth. The summary

of results (Table 21) also shows that the diet of northern

searobin from this estuary (Table 20) is consistent with other

studies (Table 21).

Striped Bass (Morone saxatilus)

Hudson-Raritan Estuary Results

Striped bass of small-to-medium size (range of 13.5-65.0

cm FL, mean of 33.2 cm) were generally only collected by

trawl within the Estuary during the fall-winter, especially in

western areas of the Estuary: Gravesend Bay (Stratum 6)

and channels (Figure 12; Wilk et al.1998). The 81 striped

bass which were examined ate a diversity of prey, with greater

than 20 identifiable species. The diet was dominated by a

variety of small or juvenile fish and crustaceans (Table 22).

Many stomachs per collection (up to 100%) were empty.

Crangon again led in the diet with an FO of 62.3% (range of

54.5-75.9%) and TV of 50.3% (range of 15.0-100.0%). All

other prey occurred or contributed less than 5% to TV, except Neomysis (Table 22).

Other prey or items found in striped bass stomachs in

lesser quantities were: polychaetes (Nephtys sp.); isopods

(Cirolana sp.); amphipods (unidentified and Gammarus

lawrencianus); mysids (Heteromysis formosa); decapod

crustaceans (Axius serratus); stomatopod (Squilla empusa);

fish (rock gunnel, juvenile searobin, juvenile unidentified

flounder, juvenile conger eel (Conger oceanicus), juvenile

Urophycis sp., northern pipefish, bay anchovy, striped anchovy (Anchoa hepsetus), juvenile winter flounder, and juvenile grubby).

Comparisons with Other Diet Studies

There are no data on the diet of striped bass from within

this part of the overall Hudson-Raritan Estuary. However,

Lawler, Matusky & Skelly Engineers (1980) and Gardinier

and Hoff (1982) did report on the striped bass diet in the

Hudson River, 50 km north of the Verrazano Narrows. There

they found that juveniles, less than 20 cm FL, fed on a mix of

freshwater and marine organisms, including Gammarus and

other amphipods (e.g., Corophium, Leptocheirus, and

Monoculodes), insect larvae, copepods, isopods (e.g.,

Cyathura sp.), polychaetes, small decapod crustaceans

(Crangon and mud crabs), and some small fish. While larger

individuals were almost totally piscivorous, preying on river

herring, Atlantic tomcod (Microgadus tomcod), bay anchovy, white perch (Morone americana), and killifish, they

occasionally ate small crustaceans. Gladden et al. (1988),

possibly summarizing the same data, also reported the species ate “fish and macro invertebrates” in the same study

area. Twenty-four, 7-39 cm FL striped bass were also collected in the Raritan River during April 1976 - March 1977,

but all of their stomachs were found empty (Lynch et al.

1977).

In general, most dietary studies of this species found

seasonal and regional variability in prey (Table 23) that often reflected differences in local environmental conditions

(e.g., salinity), in the size of the fish examined, and/or in the

time of year (e.g., Bigelow and Schroeder 1953). There is a

clear and well documented ontological shift in predation

focus from small crustaceans (e.g., copepods, amphipods,

and mysids) and small or juvenile fish for the youngest and

smallest striped bass, to larger fish and crustaceans (e.g.,

crabs and shrimp) for the older and larger striped bass. The

smaller-sized striped bass examined in the present survey

ate a mix of small crustaceans and fish (Table 22).

Clearnose Skate (Raja eglanteria)

Hudson-Raritan Estuary Results

Clearnose skate were collected in the sandier, eastern

polyhaline areas within the Estuary, such as Lower Bay,

Gravesend Bay, East Bank, Romer Shoal, Sandy Hook Bay,

and Raritan Channel (Strata 1, 3-6, 9; Figure 13), and during

the summer (Wilk et al.1998). The diet of the 71 clearnose

skate which were examined (range of 49.0-86.0 cm TL, mean

of 63.3 cm) included a diversity of crustaceans, fish, and

other prey (Table 24). Crangon, juvenile or small Atlantic

rock crabs and Ovalipes ocellatus, and fish were most frequently found in the stomachs and contributed most to

overall stomach volumes (Table 24). No empty stomachs

were found.

Other prey or items found in clearnose skate stomachs

in lesser quantities were: unidentified algae; mollusks (unidentified); amphipods (unidentified); mysids (Neomysis

americana); decapod crustaceans (Pagurus longicarpus

and juvenile Libinia sp.); and fish (unidentified juvenile

hake, juvenile striped searobin, juvenile black sea bass, rock

gunnel, juvenile searobins, and gobies).

Comparisons with Other Diet Studies

No previous studies of the diet of this species are known

for this estuary. In fact, information on the diet of this

species in general is very weak, and only available from a
Page 11

few studies (Table 25). In Delaware Bay, Fitz and Daiber

(1963) examined the diet of this species and found that it

also most commonly ate Crangon (i.e., an FO of 60%), Ensis

directus (i.e., an FO of 36%), mud crabs (i.e., an FO of 20+%),

and to a lesser degree, a variety of other small crustaceans,

bivalve mollusks, and small fish such as weakfish and windowpane. Prey volume or weight contributions were not

noted, but numerically, Crangon was still the dominant prey,

and Neomysis was second in importance. Fritz and Daiber

(1963) also noted that in the fall the skate ate more Neomysis,

decapod crustaceans, and fish, but in the spring they focused more on Crangon and Ensis. Kimmel (1973) examined a small collection of juveniles (less than 44 cm TL) of

this species at the mouth of Chesapeake Bay and found

that Crangon, Ensis, and the mud shrimp Upogebia affinis

volumetrically dominated stomach contents, but that a variety of epifaunal invertebrates (especially crustaceans) and

small fish (searobins and hake) were also eaten. The diet

described by the 1973 Kimmel paper is consistent with the

prey that Hildebrand and Schroeder (1928) noted in the few

clearnose skate that they examined from inside Chesapeake

Bay. In the present study, the only prey that was found that

have not been previously reported to be important in the

diet were Atlantic rock crabs and O. ocellatus (Tables 24

and 25).

Bluefish (Pomatomus saltatrix)

Hudson-Raritan Estuary Results

Juvenile, YOY (range of 7.0-13.5 cm FL, mean of 8.9 cm)

bluefish were collected in the summer-fall in the Estuary,

and mostly in or near channels (Figure 14; Wilk et al.1998).

The stomachs of 63 bluefish were examined; 62 of these

were from one collection – August 1997. Fish, Crangon,

and Neomysis dominated their diet (Table 26). The identifiable fish prey included mostly midwater forms: butterfish,

silversides, anchovies, but also juvenile black sea bass.

The only other prey or items found in bluefish stomachs were unidentified algae and the polychaete Nereis

succinea.

Comparisons with Other Diet Studies

Since only juvenile bluefish were collected and examined in this study, the following summary keeps that focus.

Friedland et al.(1988) examined the diet of YOY bluefish in

this estuary and reported that fish dominated the diet (by

FO and TW) in Sandy Hook Bay (Stratum 1), especially bay

anchovy, silversides, and killifish; however, Crangon were

almost equally important, along with Neomysis. Breder

(1922b) also notes that a small bluefish, also caught in Sandy

Hook Bay, had a sand lance in its stomach. A limited study

of the diet of bluefish in the Raritan River and adjacent

western Raritan Bay found that juveniles (3-22.5 cm TL)

collected by seine had eaten mummichog, bay anchovy,

silversides, Crangon, Palaemonetes sp., and unidentified

fish, while larger bluefish (greater than 37 cm FL) collected

by gill net had eaten Atlantic menhaden, spot (Leiostomus

xanthurus), bay anchovy, and Crangon (Lynch et al.1977).

 In the adjacent brackish Hudson River, YOY bluefish

consumed a variety of fish during their summer residency,

including juvenile striped bass, white perch, American shad

(Alosa sapidissima), blueback herring (A. aestivalis), Atlantic tomcod, silversides, bay anchovy, and occasionally

other species, as well as blue crabs (Juanes et al.1993; Buckel

et al.1999) (Table 27).

In nearby southern Long Island, New York, and elsewhere in the coastal Middle Atlantic Bight, juvenile bluefish were reported to commonly eat small schooling fish

such as silversides, bay anchovy, butterfish, killifishes,

juvenile Atlantic menhaden, herring, and weakfish, as well

as benthic fish such as winter flounder, spot, and Atlantic

tomcod (Table 27; Greenley 1939; Tatham et al.1984). Small

crustaceans, such as Palaemonetes sp., Crangon, and

Neomysis also dominated the YOY or juvenile bluefish diet

(Table 27). The diet of the relatively small sampling of juvenile bluefish examined in the present study (Table 26) show

that Friedland et al.’s (1988) findings are probably representative of the species’ diet within the Estuary, are typical

for this life stage of the species (Table 27), and are consistent with previous dietary summaries (Baird 1873; Nichols

and Breder 1927; Hildebrand and Schroeder 1928; Bigelow

and Schroeder 1953; Richards 1976).

Winter Skate (Raja ocellata)

Hudson-Raritan Estuary Results

Adult winter skate were collected from all areas of the

Estuary during the cooler seasons, but they were especially abundant in or near channels (Figure 15; Wilk et

al.1998). The 57 winter skate (range of 36.0-77.0 cm TL,

mean of 55.8 cm) which were examined ate a diverse diet of

benthic invertebrates and fish. Crangon was also a major

item in the diet, both in terms of FO and TV. Other crustaceans and a variety of small or juvenile fish (e.g., Atlantic

herring (Clupea harengus), sculpin, sand lance, and winter

flounder) were also commonly consumed (Table 28).

Other prey or items found in winter skate stomachs in

lesser quantities were: hydroids; nematodes (probably parasitic); gastropods (Nassarius trivittatus), bivalve mollusks

(unidentified, Mulinia lateralis, and Ensis directus); polychaetes (unidentified); mysids (Neomysis); decapod crustaceans (unidentified crab fragments, Pagurus longicarpus,

Dichelopandalus leptocerus, and juvenile blue crab); stomatopods (Squilla empusa); and fish (juvenile Atlantic her-
Page 12

ring, juvenile red hake, goby, unidentified juvenile sculpin,

and smallmouth flounder).

Comparisons with Other Diet Studies

No previous studies of the diet of this species are known

for this estuary.

Nichols and Breder (1927) and Bigelow and Schroeder

(1953) noted the importance of Atlantic rock crabs and squid

in the diet of this species in New England waters, and that

this species also ate a variety of other benthic invertebrates

(e.g., polychaetes, amphipods, shrimp, and razor clams) and

small fish, such as juvenile skates, eels, herrings, smelt,

sand lance, mackerel, butterfish, cunner, sculpins, and silver and Urophycis hake. The few available quantitative

studies, including the present study, are consistent with

this overview (Table 29), except that the present study shows

a higher use of flounder as prey (Table 28).

Black Sea Bass (Centropristis striata)

Hudson-Raritan Estuary Results

There was a significant recruitment of juvenile black

sea bass into the Estuary in the summer-fall of 1997. (Juveniles were rarely collected in other survey years, and adults

were seldom found within the Estuary.) These juveniles

were widespread in occurrence with a slight tendency to be

found in or near channels (Figure 16; Wilk et al.1998). The

August 1997 survey collected 46 juveniles – 41 with

nonempty stomachs (range of 2.9-29.0 cm TL, mean of 10.8

cm,), mostly at sites where colonies of redbeard sponge

(Microciona prolifera) were collected. Various crustaceans

dominated the diet, especially Crangon, Neomysis, and juvenile Atlantic rock crabs. The crustacean prey also included copepods, amphipods, isopods, and other small or

juvenile decapods (Table 30). Several species of small or

juvenile fish (e.g., cunner, goby, Atlantic menhaden, and

possibly anchovy) were also eaten, as were some other

benthic invertebrate taxa.

Other prey or items found in their stomachs in lesser

quantities were: poriferans (unidentified); anthozoans (unidentified); nematodes (unidentified); gastropods (juvenile

Crepidula sp.); bivalve mollusks (Ensis directus); polychaetes (unidentified and Asabellides oculata); copepods (unidentified and Pseudodiaptomus coronatus); isopods

(Edotea triloba and Cirolana concharum); amphipods

(Ericthonius sp., Stenothoe sp., and caprellids); decapod

crustaceans (juvenile Ovalipes ocellatus and Pagurus sp.);

and fish (goby, juvenile cunner, and unidentified).

Comparisons with Other Diet Studies

No previous studies of the diet of this species are known

for this estuary.

In other Middle Atlantic Bight estuaries, juvenile black

sea bass prey principally upon small benthic crustaceans

such as isopods, amphipods, small mud crabs, Crangon,

mysids, and copepods, and upon small fish such as northern pipefish, anchovies, and silversides (Hildebrand and

Schroeder 1928; Bigelow and Schroeder 1953; Richards

1963; Kimmel 1973; Allen et al.1978; Festa 1979; Orth and

Heck 1980; Werme 1981). Kimmel (1973) noted that polychaetes (e.g., Nereis sp. and Glycera sp.) can be important,

too, and that the dominant prey shifted with fish growth

(i.e., from small crustaceans such as Neomysis and various

amphipods, to decapod crabs and polychaetes).

Most of the black sea bass collected in the Estuary

were YOY and older juveniles (Wilk et al.1998), but adults in

other coastal areas have been reported to feed upon a variety of epifaunal and infaunal invertebrates, especially crustaceans, squid, and small fish (Bigelow and Schroeder 1953;

Richards 1963; Mack and Bowman 1983; Steimle and Figley

1996). The diet of the juvenile black sea bass examined in

the Estuary was dominated by small crustaceans (Table 30)

and was similar to the diet of the species reported in other

studies (Table 31).

Spot (Leiostomus xanthurus)

Hudson-Raritan Estuary Results

Spot are generally found in the Estuary in the summerfall, and were especially common in or near the Raritan Channel (Stratum 9) and Sandy Hook Bay (Stratum 1, Figure 17)

(Wilk et al.1998). Forty-seven spot (range of 12.8-18.5 cm

FL, mean of 15.4 cm) were collected in fall-winter 1996. The

tube-dwelling amphipod Ampelisca abdita dominated the

identifiable prey, but Crangon and Neomysis were also prominent. Other benthic invertebrates, including the copepod

Pseudodiaptomus coronatus, constituted the rest of the

stomach contents, which also contained a notable amount

of unidentifiable organic matter or detritus (Table 32).

Other prey or items found in spot stomachs in lesser

quantities were: green algae; bivalve mollusks (unidentified spat); polychaetes (unidentified); and amphipods

(Corophium sp., Gammarus lawrencianus, and unidentified).

Comparisons with Other Diet Studies

No previous studies of the diet of this species are known

for this estuary.

In southern New Jersey, Festa (1979) found that YOY

spot ate copepods and amphipods (e.g., Ampelisca sp.),

while larger juveniles also included a variety of polychaetes in the diet. Elsewhere, various studies show that YOY

spot ate calanoid and harpacticoid copepods, a variety of

other small crustaceans including larvae, and detritus; while

larger juveniles (11-16 cm FL) ate more amphipods such as

Ampelisca macrocephala (Table 33). Within Chesapeake
Page 13

Bay, Hildebrand and Schroeder (1928) reported that the species ate “small and minute crustaceans and annelids, together with smaller amounts of small mollusks, fish and

vegetable debris”. Smith et al.(1984) added that a wide

diversity of plant material and benthic macrofauna was eaten.

The diet of the spot examined from this estuary focused on

small benthic organisms and detritus (Table 32), and is consistent with other dietary studies for the species (Table 33).

American Lobster (Homarus americanus)

Hudson-Raritan Estuary Results

A total of 47 American lobsters were collected during

five seasons, mainly from Romer Shoal, Gravesend Bay,

Chapel Hill, and Raritan Channel (Figure 18). The collections were a mix of juveniles and small adults (range of 2.8-

9.9 cm carapace length, mean of 5.8 cm). The highly macerated state of the stomach contents, and the American

lobster’s known tendency to eat calcareous shell fragments,

make identification of all true prey tentative. Species or

items that could be identified from the diverse, particulate

material in the stomach were included in this analysis, however. The dominant items evident in the stomachs were

fragments of decapod crustaceans, especially Atlantic rock

crabs, Pagurus sp., and Ovalipes ocellatus. Other items

that were found suggest a range of taxa being eaten (i.e.,

hydroids to skate egg cases), as well as human artifacts

such as coal pebbles, fragments of plastic and rubber, and

synthetic fibers (Table 34).

Other prey or items found in American lobster stomachs in lesser quantities were: gastropods (Crepidula

fornicata, Nassarius trivittatus, Lacuna vincta, Turbonilla

sp., and Euspira operculums); bivalve mollusks (Mulinia

lateralis); polychaetes (unidentified, Nereis succinea, and

Spiochaetopterus oculatus); arachnids (juvenile Limulus

polyphemus); cirripeds (Balanus sp.); decapod crustaceans

(P. longicarpus, O. ocellatus, and juvenile Callinectes sp.);

and echinoderms (Arbacia punctulata).

Comparisons with Other Diet Studies

No previous studies of the diet of this species are known

for this estuary.

Steimle (1994) examined the diet of American lobster

collected outside the mouth of this estuary. He reported

that the diet varied among three collection sites that were

variably influenced by sewage sludge disposal, and among

bimonthly collections, although few American lobster were

collected during winter. At the least-sludge-affected sites

(probably being most appropriate for comparison with this

estuary), the American lobsters were primarily eating Atlantic rock crabs, unidentified fish, the polychaete Pherusa

affinis, and algae (Table 35). He also noted obvious human

artifacts in the stomachs, especially animal hair and synthetic fibers.

In Long Island Sound, Weiss (1970) reported American

lobsters also ate crustaceans, especially Atlantic rock crabs,

mollusks such as Lacuna vincta and the blue mussel, and

the polychaete Nereis virens. Other American lobster diet

studies have been conducted outside the Middle Atlantic

Bight area (e.g., in the sub-boreal Gulf of Maine and for

Canadian populations). The diet of small American lobsters collected in the Estuary in the present study (Table 34

) is consistent with the few available studies summarized in

Table 35, and with the more general comments of Herrick

(1911).

Tautog (Tautoga onitis)

Hudson-Raritan Estuary Results

Fifty-one tautog (range of 8.4-58.0 cm TL, mean of 37.5

cm) were collected and examined, primarily during the

warmer seasons and from Romer Shoal, East Bank,

Gravesend Bay, and, to a lesser degree, nearby areas (Figure 19). A variety of decapod crustaceans and mollusks

were the most frequently eaten prey, with Atlantic rock crabs,

xanthid crabs (including Dyspanopeus sayi), and blue mussels being prominent in the diet (Table 36).

Other prey or items found in tautog stomachs in lesser

quantities were: hydroids; gastropods (unidentified,

Crepidula sp., and unidentified eggs); bivalve mollusks

(Anadara ovalis and juvenile northern quahogs); cirripeds

(Balanus sp.); amphipods (Gammarus sp. and Ericthonius

sp.); decapod crustaceans (unidentified and juvenile Libinia

sp.); and shell hash.

Comparisons with Other Diet Studies

No previous studies of the diet of this species are known

for this estuary, although Duffy-Anderson and Able (1999)

mention that the diet of juvenile tautog held in cages in

New York harbor appears to be “harpacticoid copepods,

mysids, and amphipods.”

Steimle and Shaheen (1999) summarized the diet of tautog, which has been resummarized in this report as Table 37.

Dorf (1994) found that juveniles in Narragansett Bay, Rhode

Island, ate various amphipods and copepods (mostly

harpacticoids). Grover (1982) found a similar juvenile diet

on the ocean side of Long Island, New York, as did Sogard

(1992) in a southern New Jersey estuary. Nichols and Breder

(1927) also noted seaweed in the diet of young tautog. The

diet of older, 2-3 yr old juveniles was generally found to

shift to mollusks, primarily blue mussels (Dorf 1994; Lankford

et al.1995), but Festa (1979) reported mud crabs to be a

primary item in the diet of larger juveniles in southern New

Jersey.
Page 14

Adult tautog are generally reported to prey primarily

upon blue mussels, but also upon barnacles, crabs (Pagurus

sp., Atlantic rock, and others), sand dollars (Echinarachnius

parma), various amphipods, Crangon and other shrimp,

American lobsters, scallops and other mollusks, and polychaetes (Hildebrand and Schroeder 1928; Bigelow and

Schroeder 1953; Festa 1979; Steimle and Ogren 1982).

Steimle (in review) found that besides blue mussels, the

large anemone Metridium senile and razor clams (Ensis

directus) can be important prey in Delaware Bay. The results of the present study (Table 36) reaffirm the importance

of “shellfish,” crustaceans, and mollusks, in the tautog diet.

Smooth Dogfish (Mustelis canis)

Hudson-Raritan Estuary Results

This relatively large (range of 55.0-111.0 cm TL, mean of

74.6 cm) visitor to the Estuary was collected in modest numbers (i.e., 42 specimens) during the warm seasons of both

survey years, and mostly from Romer Shoal, East Bank,

Gravesend Bay, nearby eastern Lower Bay areas, and near

the Raritan Channel (Figure 20). It primarily ate a variety of

decapod crustaceans and mollusks, and an occasional fish.

Among the decapod prey, Crangon, Atlantic rock crabs,

and Ovalipes ocellatus were commonly eaten, and the most

notable molluscan prey was the razor clam Ensis directus

(Table 38).

Other prey or items found in smooth dogfish stomachs

in lesser quantities were: bivalve mollusks (Atlantic

surfclam); cephalopods (unidentified squid); polychaetes

(unidentified and Glycera sp.); decapod crustaceans

(Pagurus pollicaris, Pagurus sp., and Libinia sp.); stomatopods (Squilla empusa); and fish (juvenile Atlantic menhaden, northern pipefish, and lined seahorse (Hippocampus erectus)).

Comparisons with Other Diet Studies

No previous studies of the diet of this species are known

for this estuary.

Rountree and Able (1996) examined the diet of YOY of

this species in a southern New Jersey estuary and found

small Palaemonetes sp. and Crangon shrimp as the dominant prey, followed by unidentified polychaetes and crabs,

blue crabs, and a variety of other benthic invertebrates (especially crustaceans); very few fish were eaten (Table 39).

These results were similar to an early study, near Atlantic

City, New Jersey, by Breder (1921) who reported that various crabs, eel grass, detritus, and fish were the most common items in the stomachs of smooth dogfish less than 64

cm TL. Nichols and Breder (1927) noted a preference for

eating young American lobster and blue crabs, as well as

other crustaceans, fish, and a variety of benthic macrofauna.

Bigelow and Schroeder (1953) also commented on the potential heavy predation of smooth dogfish on American

lobsters in Buzzards Bay, Massachusetts, as well as predation on Atlantic menhaden and tautog. Festa (1979) examined 12 juvenile smooth dogfish from southern New Jersey

and found that blue crabs dominated (i.e., a TV of 91%) the

diet, followed by “bay” [Crangon?] and Palaemonetes

shrimp and juvenile weakfish. The present examination of

the smooth dogfish diet in the Estuary (Table 38) shows

that the species basically eats larger crustaceans and fish,

which is consistent with the results of most other studies

(Table 39).

Silver Hake (Merluccius bilinearis)

Hudson-Raritan Estuary Results

In general, silver hake were only collected as juveniles

(range of 6.5-15.0 cm TL, mean of 10.1 cm) in the Estuary,

and then primarily during the fall and in or near channels

(Figure 21; Wilk et al.1998). Juvenile silver hake were not

commonly available for examination except in November

1997 when 29 were collected. Crustaceans were the most

common and important taxa in the diet, especially Crangon,

Neomysis, Gammarus lawrencianus, and Ampelisca abdita,

but small or juvenile fish were also eaten (e.g., silver hake,

Atlantic menhaden, and probably anchovies). Both benthic

and midwater fauna were eaten (Table 40).

Other prey or items found in silver hake stomachs in

lesser quantities were: crustaceans (unidentified);

cumaceans (unidentified); isopods (Edotea triloba); amphipods (unidentified, Jassa falcata, Hippomedon serratus, and Unciola sp.); decapod crustaceans (Palaemonetes

sp.); and fish (juvenile Atlantic menhaden).

Comparisons with Other Diet Studies

No previous studies of the diet of this species are known

for this estuary.

Table 41 summarizes dietary studies for juvenile and

older silver hake that could be relevant to the present study.

Schaefer (1960) and Steimle (1985) examined stomachs of

this species collected just outside this estuary, and found

that mysids (mostly Neomysis), Crangon, small unidentifiable fish, and YOY silver hake were the most important prey

for near adult and adult fish. Schaefer (1960) also examined

adults caught by hook-and-line on a surf-zone fishing pier

in Long Branch, New Jersey (20 km south of the mouth of

the Estuary), and found a slightly different diet from that

found offshore. Inshore, he found that silver hake ate amphipods, Crangon, YOY silver hake, and mysids, in that

order of relative importance. Richards (1963) reported a
Page 15

similar diet in Long Island Sound. On the continental shelf,

Sedberry (1983) and Bowman et al.(1987) found that juvenile (i.e., less than or equal to 20 cm TL) silver hake ate

various crustaceans, including euphausids and the hyperid

amphipod Parathemisto gaudichaudi. Smaller, YOY silver

hake (i.e., less than 5 cm TL) ate benthic and pelagic amphipods; larger YOY (i.e., between 5 and 10 cm TL) ate Crangon

and Dichelopandalus pinguis [leptocerus?] shrimp, amphipods, small fish (sand lance and smaller silver hake), and

squid. The juveniles collected in the Estuary during the

present study ate a diet consistent with those studies noted

in Table 41, with such studies as Jensen and Fritz (1960) and

Vinogradov (1984), and with the generalizations of Bigelow

and Schroeder (1953). Bowman et al. (1987) report that

silver hake are mostly nocturnal feeders, and if so, mid-day

collections might involve some loss of information by the

digestion of softer prey. Few adult silver hake are collected

in the Estuary (Wilk et al.1998).

Less Abundant Predators

The following predators were collected in lesser quantities and in more limited areas. Their diets are only briefly

documented here, with identifiable prey being listed in order of their relative importance to overall stomach content

volumes.

Fourspot Flounder (Paralichthys oblongus)

Forty-one examples of this predator were collected from

several strata, and ranged between 6.7 and 33.0 cm TL (mean

of 15.1 cm). Crangon, unidentified fish, Neomysis, and unidentified decapod crustacean zoeae were prominent in the

diet.

Grubby (Myoxocephalus aenaeus)

Twenty-six specimens, ranging between 3.8 and 13.0

cm TL (mean of 7.9 cm), were collected mostly in the Lower

Bay to East Bank area (Strata 3 and 5). They ate Crangon,

Neomysis, juvenile Atlantic rock crabs, juvenile black sea

bass, the tubiculous amphipod Corophium sp., the isopod

Cyathura sp., caprellid amphipods, and the sand-tube worm

Sabellaria vulgaris.

White Perch (Morone americana)

Twenty-one white perch were collected, mostly in western Raritan Channel (Stratum 9), and ranged between 16.0

and 27.7 cm TL (mean of 21.0 cm). These fish ate Crangon,

unidentified small or juvenile fish, unidentified crustaceans,

gobies (Gobisoma sp.), Neomysis, Palaemonetes sp., and

Gammarus sp.

Northern Kingfish (Menticirrhus saxatilis)

Sixteen kingfish were collected, mostly in the Lower

Bay to East Bank area (Strata 3 and 5), and ranged between

7.5 and 16.5 cm TL (mean of 10.9 cm). They ate Crangon,

Gammarus lawrencianus, anchovies, unidentified polychaetes, unidentified crab, Neomysis, and Pagurus sp.

Smallmouth Flounder (Etropus microstomus)

Twelve specimens of this flounder were collected,

mostly in the Lower Bay to East Bank area (Strata 3 and 5),

and ranged between 11.3 and 15.0 cm TL (mean of 12.7 cm).

These fish ate Crangon, Pagurus longicarpus, Neomysis,

Pagurus sp., and Gammarus lawrencianus.

Spiny Dogfish (Squalus acanthias)

Twelve spiny dogfish were collected, mostly in the

Romer Shoal and Ambrose Channel area (Strata 4 and 7),

and ranged between 76.0 and 80.3 cm TL (mean of 77.4 cm).

They ate unidentified fish, Atlantic rock crabs, juvenile

ocean quahog, Pagurus pollicaris, Ovalipes ocellatus, and

northern pipefish.

Atlantic Tomcod (Microgadus tomcod)

Eleven Atlantic tomcod were collected in the Gravesend

and northern Lower Bay area (Strata 3 and 6), and ranged

between 7.5 cm and 9.7 cm TL (mean of 8.7 cm). They ate

Crangon and Gammarus lawrencianus.

Oyster Toadfish (Opsanus tau)

Ten toadfish were collected, mostly in Raritan Channel

(Stratum 9), and ranged between 11.5 and 24.5 cm TL (mean

of 16.1 cm). They ate Crangon, juvenile Atlantic rock crabs,

Pagurus longicarpus, and unidentified fish.

Rock Gunnel (Pholis gunnellus)

Nine samples of this species were collected in the Lower

Bay to East Bank area (Strata 3 and 5), and ranged between

5.2 and 12.3 cm TL (mean of 9.4 cm). They ate Neomysis,

Photis sp., unidentified isopods, Leptocheirus pinguis, and

unidentified copepods.
Page 16

Cunner (Tautogolabrus adspersus)

Nine cunner were collected, mostly in the Sandy Hook

Bay to East Bank area (Strata 4 and 5), and ranged between

3.2 and 12.2 cm TL (mean of 5.5 cm). Unidentified amphipods, harpacticoid copepods, Gammarus lawrencianus,

Neomysis, Corophium sp., Ampelisca abdita, Ericthonius

sp., unidentified foraminifera, and Unciola sp. were found

in their stomachs.

Northern Puffer (Sphoeroides maculatus)

Eight puffer were collected in Sandy Hook Bay (Stratum 1), and ranged between 7.3 and 16.2 cm TL (mean of

10.0 cm). They ate the sand-tube worm Sabellaria vulgaris, Atlantic rock crabs, unidentified crabs, hydroids,

Pagurus longicarpus, Ampelisca abdita, gastropod eggs,

barnacles, algae, and wood fragments.

Atlantic Croaker (Micropogonias undulatus)

Four croaker were collected from Raritan Channel (Stratum 9), and ranged between 12.6 and 18.0 cm TL (mean of

15.6 cm). They ate Crangon, an unidentified clam, Ampelisca

abdita, Neomysis, Glycera sp., and unidentified fish.

Longhorn Sculpin (Myoxocephalus octodecemspinosus)

Three specimens were collected from the East BankAmbrose Channel area (Strata 5 and 7), and ranged in length

between 9.0 and 29.0 cm TL (mean of 22.2 cm). They ate

sand lance, Crangon, unidentified fish, and juvenile Atlantic rock crabs.

Conger Eel (Conger oceanicus)

Three juvenile congers were collected (i.e., shaken out

of discarded beverage containers brought up in the trawl)

in Gravesend Bay (Stratum 6), and ranged between 20.5 and

30.2 cm TL (mean of 24.6 cm). They ate Pagurus longicarpus,

juvenile Atlantic rock crabs, Crangon, and mud (xanthid)

crabs.

Overall Perspective on Diets

The predator collection discussed in the preceding text

appears representative of what is typically found within the

Estuary. Wilk et al.(1998) listed 17 of the 20 predators examined in this dietary study as being among the most commonly collected species within the Estuary. The other species that they found to be common were pelagic or “forage”

species such as bay anchovy, herrings, butterfish, and

longfin inshore squid (Loligo pealeii) which are discussed

subsequently. Three predators that were examined in this

study, but that were not listed as the most common in the

trawl survey, were smooth dogfish, tautog, and American

lobster. These species were examined either because of

their fishery importance (tautog and American lobster) or

because of their being among the largest apex predators

found within the Estuary (smooth dogfish).

The species occurring in the Estuary in the 1990s appear to be persistent since 1970s, i.e., the dominant species

were consistent with those reported by Wilk and Silverman

(1976). The fish community defined by Wilk et al.(1998) in

this estuary is also similar, with a difference of only one or

two dominant species, to that found in other larger Middle

Atlantic Bight estuaries, e.g., Narragansett Bay, Rhode Island (Oviatt and Nixon 1973), Long Island Sound (Richards

1963), and Delaware Bay (Grimes 1983). However, as prey

availability may differ in those estuaries, the diets discussed

above may not adequately represent the situation for other

estuaries.

Examination of diets from trawl-collected fish can involve biases related to the collection method. Some prey

that were fresh and readily identified in the stomachs can

be an artifact of within-trawl predation. This potential bias

exists in most diet data based on trawl-caught samples, and

is likely to involve the use of larger or motile epibenthic

prey such as small fish, Crangon, and crabs that accumulate within the trawl’s cod-end, or that are disturbed by the

trawl doors, warps, footrope, or tickler chain to expose them

to rapid-response predation. These results are also subject

to other potential biases or errors that are typical of the

method. For example, differential rates of prey digestion

and stomach evacuation can be a bias, especially for afternoon collections (assuming diurnal predators often feed

heavily in the morning), although American lobsters and

other predators such as red hake are thought to be primarily

nocturnal feeders.

FORAGE BASE

Examination of the diets of common predators provides

insight into the value of various estuarine prey and habitats to support fishery populations. Also, the conservation of the habitat of prey that can be essential to fishery

resources is a requirement of the Magnuson-Stevens Fishery Conservation and Management Act (October 1996).

Consequently, for those prey which were examined in this

study and which were found to be eaten more commonly

than others, a brief overview of their life histories and a

discussion of their habitat use are presented to facilitate

effective habitat management. The following section summarizes what is known of the life histories and habitat use
Page 17

of both the commonly eaten invertebrate prey as well as

those fish that are less important to diets, but can be of

interest to fishery management.

The prey that seem to be most widely used or to be

eaten at the highest frequency or volume levels by one or

more of the predators covered in the previous sections are

summarized in Tables 42-44; Tables 42 and 43 list nonfish

used as prey, and Table 44 lists fish used as prey. These

prey are listed in order of their overall importance as prey

within the Estuary, based on the ranking of their percent

frequency of occurrence in the diet or percent contribution

to total stomach content volume, relative to the predators

examined in this study.

Dominant Invertebrate Prey and Their Life

Histories and Habitats

Sevenspine Bay Shrimp (Crangon septemspinosa)

The epibenthic, sevenspine bay shrimp (or sand shrimp)

ranked first in importance to overall diet volumes for most

of the predators. It was the most common prey of little

skate, juvenile summer flounder, juvenile red hake, weakfish, spotted hake, striped and northern searobins, juvenile

striped bass, clearnose and winter skates, juvenile black

sea bass, silver hake, and fourspot flounder (Tables 6, 10,

12, 14, 16, 18, 20, 24, 28, 30, and 40). Only winter flounder

and tautog did not rely heavily on Crangon as prey, although it was occasionally eaten by both.

Crangon occurs on sandy to silty-sand sediments into

which it can partially bury. It tolerates a wide range of

salinity and temperatures, and occurs within estuaries and

bays, offshore to about 90 m in depth, and from the subArctic to Florida (Caracciolo and Steimle 1983). It is considered omnivorous, and will eat detritus, small invertebrates,

and newly settled, postlarval fish such as flounder (Witting

and Able 1993). It breeds throughout the warmer months,

with prominent spring and weaker fall peaks known for some

areas (Wehrtmann 1994). It can live for 2 yr and grow to

about 7 cm TL.

Because Crangon is relatively motile and small, its abundance and distribution within the Estuary are not accurately known, although an unsuccessful effort was made to

obtain such information (R. Reid, unpubl. data, National

Marine Fisheries Serv., Highlands, NJ). It can avoid benthic

grab samplers and pass through the mesh of standard trawls.

Because of its importance to the diets of most fishery resources found in the Estuary, more should be known of its

preferred habitats and sensitivity to human perturbation,

including toxic chemical contaminant bioaccumulation. The

species was also once found to be affected by “black spot

disease,” or chitinoclasia, within this estuary (Gopalan and

Young 1975). The toxicity of chemically contaminated sediments in the Estuary to several crustacean species has been

reported by Long et al.(1995), but tests were not conducted

with Crangon, although tests on Palaemonetes pugio, a

marsh dweller, were mentioned.

Neomysis americana

 This mysid was generally second in overall importance,

volumetrically, to diets. It was ranked first as prey for windowpane and second in overall importance to a variety of

predators that focused upon Crangon as prey, except

skates (Table 42). It was not found or identified in the diets

of American lobster, tautog, and smooth dogfish in the size

ranges examined in the present study.

Neomysis is a dominant component of the suprabenthic/

planktonic community in most Middle Atlantic Bight estuarine ecosystems, but occurs widely along the Western Atlantic coast from Nova Scotia to the Caribbean Sea.

Caracciolo and Steimle (1983) and Hargreaves (1995) report

that it tolerates a wide range of salinities (i.e., from marine to

as low as 1‰) and temperatures (i.e., from 0 to 25°C), and

prefers to be over sandy sediment in depths less than 60 m.

It occurs in swarms that are negatively phototaxic, and

avoids strong light. It is omnivorous, eating mainly

microalgae, zooplankton, and organic microdetritus. The

abundance and distribution of Neomysis in the Estuary are

unknown at present, but the species’ photophobic nature

suggests that deep channels and depressions in the Estuary with sandy sediments may be important daylight habitat, and when they may be most available as prey to demersal predators. As for Crangon, more needs to be known on

Neomysis’s distribution, habitat use, and sensitivity to human perturbations in this estuary.

Gammarus lawrencianus

This semipelagic amphipod (also called “scud”) ranked

third in overall importance, being found in 68% of the diets

examined. It was especially important as prey to windowpane, juvenile scup, juvenile red hake, juvenile weakfish,

spotted hake, northern searobin, juvenile silver hake, and

northern kingfish (Table 42).

G. lawrencianus was reported by Bousfield (1973) to

occur on or over sandy or muddy areas of estuarine areas

of Southern New England. Amphipods of the genus

Gammarus typically move across the bottom on their sides,

and their laterally compressed body allows them to move

readily within cracks and spaces among algae and other

objects. At summer breeding times, a species of Gammarus

was observed swarming in the evening at the water surface

within the Estuary (Grant 1984), suggesting a period of enhanced exposure to predation. Little is known of the relative abundance and distribution of Gammarus in this estuary, but the species’ association with vegetation and other
Page 18

shelter, as well as its semipelagic habits, make it difficult to

survey. Sage and Herman (1972), in a rare reference to the

genus in this estuary, found that G. fasciatus was only

common in Sandy Hook Bay (Stratum 1) during November.

Atlantic Rock Crab (Cancer irroratus)

Juvenile or small Atlantic rock crabs were eaten by 60%

of the predators, being most important to larger species

such as little skate, striped searobin, summer flounder,

clearnose skate, winter skate, American lobster, tautog,

smooth dogfish, and to some juveniles such as those of

black sea bass (Table 42). This prey was usually eaten in its

YOY stage during the summer-fall, but soft-shelled stages

of larger crabs were also eaten at all seasons.

The postmegalop, juvenile stages of this crab generally appear as part of the benthic macrofauna in early summer (Steimle and Stone 1973), and this coincides with their

appearance in the diets of juvenile fish using this estuary

(e.g., winter flounder, scup, summer flounder, and black sea

bass; Tables 2, 8, 10, 30) and in the diets of other small

predators such as northern searobin (Table 20). Larger juvenile and small adult Atlantic rock crabs are eaten by other

predators during other seasons; these larger crabs appear

to leave this estuary in summer, except in deeper channels

(Wilk et al.1998). Studies of this crab suggest that, despite

its common name, “rock crab,” it is more commonly found

on sand bottoms than on gravel and rock (e.g., Jeffries 1966;

Bigford 1979; Palma et al.1999). However, the use of rough

bottom or rock habitats by motile invertebrates and fish is

poorly known because of sampling problems and inadequate

survey effort (Steimle and Zetlin 2000). Reilly and Saila

(1978) used diver surveys and reported mussel beds as a

preferred habitat for juvenile Atlantic rock crabs off Southern New England. Atlantic rock crabs were most commonly

collected by trawl in and near channels and throughout the

marine, eastern part of the Estuary (Figure 22).

Lady Crab (Ovalipes ocellatus)

The lady or calico crab is a warm-season (April-December) ecosystem component, being collected from all areas

and strata (Figure 23), although it is reported to prefer sands

(Stehlik et al.1991). Juveniles of this species were also

most often found in diets. YOY seem to be available as prey

in the summer and fall in this estuary, but larger (i.e., 20-30

mm) crabs were collected in the spring too (L. Stehlik, pers.

comm., National Marine Fisheries Serv., Highlands, NJ).

Right-Handed Hermit Crabs (Pagurus spp.)

Several species of hermit crabs (Pagurus spp.) were

eaten by predators (i.e., an overall FO of 64%), but these

prey seemed only really important to American lobster and

smallmouth flounder (Table 42). Usually only the distinct,

distal ends of the legs and claws were identifiable in lobster

stomachs. Two species were readily identified as prey, the

small P. longicarpus and the larger P. pollicaris, the latter

of which was rarely found as prey. Both species are considered omnivors/detritivors and are reported to be common

on a wide range of habitats. P. longicarpus is found in

shallow waters in the summer (including intertidal and lower

salinity areas) from Canada to Texas, but migrates to deeper

water and sandy bottoms as waters cool, where it often

hibernates in pits that it digs (Rebach 1974). P. pollicaris

tends to stay in deeper, more saline waters with sandy sediments.

Ampelisca abdita

This tube-dwelling amphipod was next in overall dietary importance, occurring in the diets of 56% of the predators examined. It was particularly important to the diets of

winter flounder, windowpane, juvenile scup, juvenile weakfish, striped searobins, juvenile black sea bass, and juvenile silver hake (Table 42). Recent benthic surveys of this

estuary (Cerrato et al.1989) found this species to be common throughout the year in silty areas such as Sandy Hook

Bay (Stratum 1), in western areas of the Estuary (Strata 2

and 9), and in Gravesend Bay (Stratum 6). Considering this

distribution, it is curious that Long et al.(1995) commented

on tests of the toxicity of sediment from various locations

within the Estuary to this species, and noted that, in the

1980s and early 1990s, western Raritan Bay silty sediments

(western parts of Strata 2, 3, and 9) were found to be toxic,

while sediments from sandy areas in the northeastern third

of the Estuary (eastern part of Stratum 3, and Strata 4, 5, 7,

and 8) were relatively low in toxicity. Wide variance in annual abundances have been reported for this species

(Steimle and Caracciolo-Ward 1989).

Northern Quahog (Mercenaria mercenaria) and Atlantic

Surfclam (Spisula solidissima) Siphons

This type of prey was important only to winter flounder, and was eaten infrequently by other predators (Table

42). Because winter flounder and these two clams are dominant and fishery important species within the Estuary, the

clams are included in this discussion. Northern quahogs

are generally found in the fine-sand, silty central, western,

and southern areas of the Estuary (de Falco 1967; McCloy

1984; Cerrato et al.1989), basically in Strata 1-3 and 9, and in

the deep silty area in Gravesend Bay (Stratum 6). Atlantic

surfclams occur in the eastern, marine areas of the Estuary

(Cerrato et al.1989), i.e., Strata 4-7. Thus, either one or the

other of these two species of larger clams is available

throughout the Estuary for siphon predation by winter flounder. The use of siphons seems unrelated to the availability

of polychaetes and amphipods as potential prey, based on
Page 19

the results of Steimle and Caracciolo-Ward (1989) and

Cerrato et al.(1989), who show that these taxa were generally available, although at lesser biomass levels during the

winter, and at quantities comparable to those in other estuaries (Steimle and Caracciolo-Ward 1989). Predation on clam

tissue would appear not to have any energetic advantage,

as clam tissue has one-half to one-third of the caloric foodenergy value of polychaetes or crustaceans (Steimle and

Terranova 1985), and tearing off a piece of relatively tough

and muscular siphon must entail more effort than picking

up unattached prey off the sediment surface. Thus, this

focused use of siphons seems to be an enigma, although it

could be related to a declining supply of benthos in the fall,

as is typically found in many estuaries and coastal areas

(Steimle 1985, 1990). Brief habitat summaries of other, lessused prey are presented in Table 43.

Habitat and Community Associations of

Invertebrate Prey

Habitat Associations

The invertebrate prey discussed in the preceding section come from one of three general habitat-associated

groups within the Estuary: endobenthic, epibenthic, and

suprabenthic (semipelagic). The endobenthic (or infaunal)

prey group includes organisms living within the sediment

or in tubes upon the sediment surface, and consists primarily of mollusks (often only their siphons being eaten), polychaetes, and certain amphipods such as Ampelisca abdita,

Corophium sp., and Unciola sp. (Table 42). This prey group

generally uses sediment carbon (including bacteria and

meiofauna) or surficial phytoplankton as food. This group

is exploited as prey by a limited group of predators: winter

flounder, scup, and spot (Table 42).

The epibenthic prey group consists of mostly motile

species, especially small decapod crustaceans that move

slowly across the sediment surface (e.g., hermit, Atlantic

rock, lady, and other crabs, and Crangon; Table 42). These

prey tend to be omnivores, capable of using detritus as well

as smaller organisms they encounter on the bottom, including larval fish (Witting and Able 1993). This group, especially Crangon, is exploited by the widest range of predators in this estuary.

The suprabenthic, or semipelagic, prey group consists

primarily of Neomysis, seasonally augmented by gammarid

amphipods such as Gammarus annulatus or G.

lawrencianus, and by copepods that may be abundant near

the bottom (e.g., Pseudodiaptomus coronatus). These prey

typically occur in swarms, and Neomysis can spend the

daytime close to the bottom, but move up in the water column at night. They tend to be planktivors, although the

gammarids may be capable of exploiting a wider range of

small food items, including the scavenging of carrion.

The high use of crustaceans as prey in all three of these

habitat-associated groups seems to be typical of food webs

in many estuaries, and has apparently not been significantly

altered for decades in the Estuary, e.g., see Townes (1939).

Townes (1939) also concludes that crustaceans are the most

important prey of fish in New York coastal waters. He noted

that “shrimp” (Crangon and Palaemonetes) and “opossum

shrimp” (Neomysis) were the most important prey, but amphipods and other taxa were important, too.

Community Associations

Frame (1974), MacPhee (1969), and others have commented that the food of marine predators generally reflects

the environmental conditions and habitats in which the

predators live. Following is a brief review of where the

aforementioned invertebrate prey can be expected to be

found within the Estuary, based on available survey information. Because of logistic constraints, benthic invertebrate collections were not a feature of the 1996-97 survey.

However, there have been several recent studies of the

benthic community in the Estuary which have characterized the major community types and their distributions

(Cerrato et al.1989; Steimle and Caracciolo-Ward 1989;

Wilber et al., unpubl. data, National Ocean Serv., Charleston, SC).

Benthic organisms are known to exhibit wide variances

in abundance, especially the smaller, short-lived species,

but benthic communities, in general, are consistently associated with certain sediment characteristics and thus can be

conservative over time, even if many community members

fluctuate in abundance. Steimle and Caracciolo-Ward (1989)

examined the information available on the Estuary’s benthic

community to the mid-1970s, Cerrato et al.(1989) examined

the fauna in the mid-1980s, and Wilber et al.(unpubl. data,

National Ocean Serv., Charleston, SC) examined the benthic

fauna in the mid-1990s. All of these studies report both

similarities and differences of major community types within

the Estuary (as defined by dominant species), that are sediment and water depth related, for the most part.

Steimle and Caracciolo-Ward (1989) defined a silty sediment community numerically dominated by several species

of polychaetes (e.g., spionids, Nephtys picta, and Sabellaria

vulgaris), mollusks (e.g., Mulinia lateralis, Acteon

punctostratus, Tellina agilis, and Nassarius trivittatus),

and a few amphipods (e.g., Rhepoxynius epistomus). Examination of biomass data identified: 1) Nephtys incisa as

important in muddy areas (such as Strata 1-2 and 6); 2)

Glycera sp., Nassarius trivittatus, and Tellina agilis as important in Lower Bay sands (Strata 3 and 4); 3) Mulinia

lateralis as important in Sandy Hook Bay (Stratum 1); and

4) Crangon, Pagurus sp., and Dyspanopeus sayi as important in scattered areas. These authors noted that this community may have been stressed from a severe tropical storm

(hurricane Agnes) that passed through the area during the

previous year.

Cerrato et al. (1989) found a different mix of common

species in the same areas reported by Steimle and Caracciolo-
Page 20

Ward (1989). They noted dominant species and their general seasonal and spatial distributions; a majority of the

benthic infaunal species that they found to be common in

the mid-1980s were found to be numerically common prey

items in the diets examined in the present study: Ampelisca

abdita, Asabellides oculata, blue mussel spat, Crepidula

fornicata, Corophium tuberculatum, northern quahogs,

and Atlantic surfclams. This similarity of common infauna

found by Cerrato et al.(1989) and common prey found in the

present study suggests that the benthic community found

in the mid-1980s persisted to the mid-1990 period of the diet

survey. These benthic studies all reported that, overall, the

benthic invertebrates within the Estuary were most abundant in a silty band that occurred: 1) from Sandy Hook Bay

northwest to off Princes Bay, Staten Island (i.e., basically

Stratum 1 and the deeper areas of Strata 2 and 3); 2) adjacent to Raritan Channel (Stratum 9); and 3) in areas of

Gravesend Bay (Stratum 6). The species which were most

abundant in this band included those which were prey, such

as Ampelisca abdita, Asabellides oculata, C. tuberculatum,

and northern quahogs. Overall, benthic invertebrates were

least abundant in the eastern, fine-to-medium sand habitats

of Strata 4 and 5, and the eastern parts of Stratum 3; the

species found to be most common in these areas were blue

mussel spat and Atlantic surfclams, which are also prey

species.

The most recent benthic survey of the Estuary, during

October 1994 and June 1995 (Wilber et al., unpubl. data,

National Ocean Serv., Charleston, SC), noted the distribution of several benthic species that were primary prey in the

present study; but their survey did not sample the channels (Strata 7-9). Atlantic surfclams and northern quahogs,

the siphons of which were a major prey type for winter

flounder, were most commonly collected in two separate

areas of the Estuary: Atlantic surfclams within sandy Strata

4 and 5 and the northeastern third of Stratum 3, and northern quahogs in the silty parts of Strata 1 and 2 and lower

half of Stratum 3. Blue mussels, a less common prey of

several species, were collected in scattered locations across

the eastern, sandy half of the Estuary, including near banks,

shoals, and former borrow pits, in Strata 3-6. Blue mussels

were especially common in June 1995, suggesting a strong

spring 1997 recruitment occurred. The amphipod Ampelisca

abdita was collected widely within the Estuary during both

sampling periods, but especially in the silty parts of Strata

1-3 and 6, and irregularly in Strata 4 and 5.

The results of the most recent benthic studies by

Cerrato et al. (1989) and Wilber et al. (unpubl. data, National Ocean Serv., Charleston, SC) are consistent, and suggest that certain recent benthic prey communities have persisted in their relative abundance and distribution within

the Estuary. Their results are most likely representative of

the prey community that would be available during this

1996-97 diet study, especially of those species that are relatively long lived such as the clams. There were, however,

also benthic invertebrates they defined as abundant that

did not appear in the present study to be readily used as

prey: the minute capitellid “thread” worms Heteromastis

filiformis and Mediomastus spp.; the spionid polychaete

Streplospio benedicti; and the small bivalve mollusks

softshell, Tellina agilis, Macoma baltica, Gemma gemma,

and Mulinia lateralis. Steimle et al.(1994) found that winter flounder collected outside the Estuary often, but not

always, ate what was most available (i.e., abundant and of a

suitable size) in the benthic community.

Fish as Prey and Their Life Histories and Habitats

Fish are also eaten by a number of predators in the

Estuary, especially by summer flounder, weakfish, spotted

and silver hakes, striped bass, bluefish, and clearnose and

winter skates (Tables 10, 14, 16, 22, 24, 26, 28, and 40). The

bay anchovy seems to be a favorite prey for many predators, but a number of demersal or benthic fish species are

eaten frequently, too. It seems few juvenile or small fish are

relatively immune from predation. None of the piscivors

seem to be obligatory and almost all rely on epibenthic

invertebrate prey to a high extent. Below is a summary of

the fish observed in their stomachs, and some brief notes

on habitats known to be commonly used by this prey group.

Twenty-six species of fish were identified in the stomachs of the predators examined in this study; other fish

remains could not be identified to the species level (Table

44). Fish, at some level of identification, occurred in the

diets of all but three predators, these exceptions being winter flounder, spot, and tautog. No specific fish species was

widely eaten by predators. The maximum FO of prey among

predators was 28% for anchovy being eaten by smallmouth

flounder. Most fish that were found in the stomachs were

juveniles, but some were mixed sizes, including adults of

such small species as rock gunnel, northern searobin, smallmouth flounder, goby, northern pipefish, and lined seahorse.

Most fishes eaten by predators were demersal, except anchovies, Atlantic menhaden, Atlantic herring, Alosa herring, Menidia sp., juvenile weakfish, butterfish, and possibly juvenile silver hake. Most fishes found in the Estuary

seem to be preyed upon by larger fish, except elasmobranchs

(i.e., skates and dogfish sharks), although fragments of skate

egg cases (possibly empty) were found in some American

lobster stomachs.

Following are some brief notes on the life history and

habitat use of those fish most commonly eaten as prey,

presented as a convenient reference for habitat managers

involved in this estuary.

Searobins (Prionotus spp.)

All but the adults of the striped searobin were eaten by

a number of predators, including incidences of cannibalism

(Table 44). Able and Fahay (1998) reported that juveniles of
Page 21

both species prefer sandy sediments when they occur in

estuaries during the late spring to fall, although northern

searobin will move out of heated shallow waters to deeper,

cooler water during mid-summer. Both species winter offshore.

Bay Anchovy (Anchoa mitchilli)

This is a common nektonic species and important prey

for a variety of fish in this and other Middle Atlantic Bight

estuaries (Houde and Zastrow 1991; Wilk et al.1998). It is a

small, schooling species that feeds on zooplankton (including fish and decapod crustacean eggs and larvae), and tolerates a wide range of salinity and temperature, although its

abundance and distribution within estuaries can vary annually (Houde and Zastrow 1991). It grows to about 9 cm

TL, and can live for about 3 yr. Spawning begins at age 1

and occurs in the spring and summer. The eggs hatch in

about 1 day, and the larvae and early juveniles are common

in the summer/fall and can be an important prey for juvenile

weakfish (e.g., Richards (1963), Van Engle and Joseph (1968),

and others (Table 44)) and many other predators found in

the Estuary. Wilk et al.(1998) show that, within the Estuary,

bay anchovy was most frequently collected by trawls in

Strata 1-3 (near Staten Island), 6, and channel Strata 8 and 9

(Figure 24), but were not found in the winter within this

estuary.

Silver Hake (Merluccius bilinearis)

Although juveniles are commonly found within the Estuary during the cooler months (Wilk et al.1998), all life

stages of this species were reported rare within central New

Jersey estuaries (Able and Fahay 1998). This species is

usually considered nektonic, but Auster et al. (1997) reported that juveniles of this species occur at high densities

in patches of amphipod tubes and other complex, structured benthic habitats found on the continental shelf.

Cunner (Tautogolabrus adspersus)

This small species is usually found year-round in marine parts of estuaries, and associated with structure such

as shellfish and seagrass/algal beds, piers, pilings, bridge

abutments, rock rip-rap, etc., although it can be less common in the winter (Able and Fahay 1998). In the trawl survey of Wilk et al. (1998), smaller sizes of cunner, which

usually occur in stomachs, were usually collected when the

trawl picked up some large debris (such as automobile tires),

snagged on some object on the bottom, or caught a quantity of redbeard sponge. This reliance on shelter and rough

bottom, which interferes with trawling, can explain why the

species has been reported to be rare in previous trawl surveys of the Estuary (Berg and Levinton 1985). Its occurrence in the stomachs of open-bottom predators, such as

summer flounder and skate (Table 44), suggests that cunner can be available away from, but probably near, shelter at

times.

Silverside (Menidia sp.)

This common “baitfish” is usually found over shallow,

sandy, nearshore areas, but was only identified in juvenile

bluefish stomachs (Table 44). Conover and Murawski (1982)

report that the species will move out of estuaries during the

fall-winter in the northern part of its range.

Winter Flounder (Pseudopleuronectes americanus)

Although not normally considered a prey species, juvenile winter flounder were relatively common in skate diets in this estuary (Table 44), and Manderson et al.(1999)

found that a limited sampling of adult striped searobins

suggested that juvenile winter flounder were preyed upon

in shallow water of Sandy Hook Bay (Stratum 1). Many of

the unidentified small flounder remains found in the stomachs examined in the present study (Table 44) may also be

this species, although both windowpane and smallmouth

flounder are found in this size range. This is a well studied,

estuarine-coastal species (Pereira et al.1999), and its life

history and habitat uses need not be reiterated here.

Sand Lance (Ammodytes sp.)

Able and Fahay (1998) reported that this species is common in shallow marine waters where it occurs in schools

that often dive and burrow into sands when threatened.

Skates appear to be its primary predator within the Estuary

(Table 44). Sand lance were rarely collected in the recent

trawl surveys within the Estuary (e.g., see Wilk et al.1998).

Butterfish (Peprilus triacanthus)

Able and Fahay (1998) summarized the life history and

habitat use of this pelagic species which is found within

estuaries during the summer, including the Hudson-Raritan

(Wilk et al.1998). This species was collected from all survey

strata, although collections were relatively scarce within

Sandy Hook Bay (Stratum 1). Able and Fahay (1998) noted

that this species is negatively phototaxic and stays near

the bottom during the day, making it most accessible to

demersal predators at that time. However, only the juveniles of bluefish and weakfish seem to eat it (Table 44).
Page 22

Early juveniles have been reported to be associated with

and to accompany jellyfish when they appear inshore during the summer (Bigelow and Schroeder 1953).

American Eel (Anguilla rostrata)

Juveniles (elvers and “whips”) of this eel were found in

the stomachs of skate and adult spotted hake (Table 44).

Able and Fahay (1998) reported that the juvenile and other

phases of this species are common in estuaries, especially

the Hudson River, where they occur in a wide range of habitats, silty to sandy, structured to unstructured.

Overall Perspective on Forage Base

Overall, the fish that were eaten as prey in this estuary

were not as important as crustaceans and some other invertebrates as food for the bulk of the fish found in the Estuary. However, fish were a common prey type for some predators, and at times could have a notable impact on the survival and recruitment of some estuarine-dependent species.

Understanding this relationship and its potential is important in developing a better understanding of the role of the

benthic environment in fishery resource production.

Information on the diets of common fish and American

lobster collected in the stressed Hudson-Raritan Estuary

shows that these diets were still similar to the diets of the

same species in other larger, less-stressed Middle Atlantic

Bight estuaries. This suggests that the Estuary still functions tropho-dynamically like other, less-stressed areas. The

study results also suggest that the Estuary’s benthic habitat and macrofauna community are important to support a

sustainable, multispecies fishery (and other biological resources such as wintering, fish-eating ducks). Steimle and

Caracciolo-Ward (1989) and recent surveys all suggest that

the Estuary’s benthic fauna seem to be relatively healthy

compared with other major estuaries, and that most of its

expected community structure is still intact, although longterm trend data are absent. There could still be a potential

problem with contamination of benthic prey by toxic substances from the Estuary’s sediments and water, and from

biotransfer up the food web, as suggested by Long et al.

(1995).

ACKNOWLEDGMENTS

 We thank Barbara Lamp for supplying New Jersey

Marine Academy of Science and Technology and other

student volunteers, including Andy Craig. We also thank

colleagues Eileen MacHaffie, Steve Fromm, Joe Vitaliano,

and Bob Reid. The cooperation of the crew of the NOAA

R/V Gloria Michelle (Eric Nelson, Jeff Broeneck, and Jeremy Adams), Sally Swarts of UNICOR-ADP, and Sue Fromm

are also recognized. Mike Fahay assisted in the identification of juvenile fish, and Pat Shaheen provided support on

copepod taxonomy. We also thank Jason Link and others

for their constructive comments.

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Williams, A.B.; Abele,.L.G.; Felder, D.L.; Hobbs, H.H.; Manning, R.B.; McLaughlin, P.A.; Perez Farfante, I. 1988.

Common and scientific names of aquatic invertebrates

from the United States and Canada: decapod crustaceans.

Am. Fish. Soc. Spec. Publ. 17; 77 p.

Witting, D.A.; Able, K.W. 1993. Effects of body size on probability of predation for juvenile summer and winter flounder based on laboratory experiments. Fish. Bull. (Wash.,

D.C.) 91:577-581.

Wolfe, D.A.; Long, E.R.; Thursby, G.B. 1996. Sediment toxicity in the Hudson-Raritan Estuary: distribution and correlations with chemical contamination. Estuaries 19:901-

912.

Worobec, M.N. 1984. Field estimate of daily ration of winter

flounder, Pseudopleuronectes americanus (Walbaum) in

a Southern New England salt pond. J. Exp. Mar. Biol.

Ecol. 77:183-196.
Page 28

Table 1. Habitat characteristics of strata sampled in the Hudson-Raritan Estuary, 1992-97. (See Figure 1 for boundaries and areas of each stratum.)

 Salinity Range Dissolved Oxygen Temperature Habitat Types/ Dominant Benthic

Stratum Depths (m)a Sedimentsb (ppt)a Range (mg/l)a Range (°C)a Currentsc Structure Community Typesb

1- Sandy Hook 3.5-10.0 (avg. = >50% silt-clay, except ~15-27 2.0-13.0; avg. = 0.3-23.8; avg. = ~<0.6 knots; Protected. Gradual Numerically abundant

Bay 6.2); deepest to nearshore. Moderate to 8.7 11.1 north-south (<1%) slope, except infauna, often

east and off Sandy high chemical (toxic abrupt shoals parallel including Ampelisca

Hook Point metals and organics) to Sandy Hook. Ulva, abdita, northern

contamination. other algae, redbeard quahogs, softshells,

sponge, and terrestrial and blue mussels.plant debris commonin fall.

2- Raritan Bay 3.3-14.0 (avg. = >50% silt-clay in and ~16-26; lowest 3.6-13.2; avg. = 0.0-24.8; avg. = ~<0.7 knots; Semiprotected. Flat, Patchy abundance,

6.6); deepest near west of Keyport Harbor to southwest 8.9 11.0 east-west gradual (<1%) slope; lowest to west.Raritan Channel and near Earle terminal the 3-m trawl depth Ampelisca abdita

channel; gravelly coarse limit excludes much of often common insand to silty sand in the wide shallow area deeper, siltier areasbetween and in off the New Jersey near Raritanshallower areas. coast, especially to the Channel; patchesModerate to high west; Raritan Channel is of Atlantic surfclam,chemical contamination along the north northern quahog, andin silty areas. boundary. softshell abundance.

3- Lower Bay 3.3-13.8 (avg. = Sand, except mud in ~16-29; lowest 5.1-13.8; avg. = 0.0-26.1; avg. ~<0.7 knots, Partially exposed. Overall numerical

6.6); deepest in borrow pits and deepest to west 8.9 = 11.1 except near Gradual (<1%) slope, abundance variable,West Bank borrow areas near Raritan Chapel Hill interrupted by borrow lowest on flats offpits and near Channel and west of Channel; east- pits north and south of New Dorp Beach.

channels Old Orchard Shoal. west, but north- West Bank, bordered by Ampelisca abdita,

Moderate chemical south near channels south and east. other amphipods,contamination in silty West Bank Sponge patches in bivalve mollusks,areas. western areas. and polychaetes

abundant in deeper,siltier areas alongsouthern boundary.

4- Romer Shoal, 3.5-9.0 (avg. = Coarse to medium ~15-30 5.0-12.4; avg. = 0.9-23.7; avg. ~0.4->2.5 Exposed to the sea. Overall abundanceFlynns Knoll, 7.0); deepest in sand with shell. Low 8.8 = 11.1 knots, generally Two shoals split by highest on RomerSwash Channel Swash Channel. sediment chemical northwest- the Swash Channel, Shoal, <1000 ind./ m2

contamination. southeast and bordered by three elsewhere. Blue

east-west channels. Blue mussel mussels and Atlantic

beds are common. surfclams common.

5- East Bank 4.0-22.0 (avg. = Coarse to medium ~20-31 5.0-12.4; avg. = 0.6-23.0; avg. ~0.2-2.0 Exposed to the sea. Overall abundance

7.5); deepest near sand, and gravelly sand 8.8 = 11.2 knots, generally Complex bathymetry highest near BreezyAmbrose Channel with shell near Breezy northwest- including shoals and and Norton Pointsand seaward of Point. Low chemical southeast natural and dredged because of blue

East Bank contamination. channels. Blue mussel mussels. Atlantic

beds are common. surfclams common

elsewhere.
Page 29

6- Gravesend 4.3-26.0 (below >50% silt near the ~19-29 5.0-12.0; avg. = 0.2-22.9; avg. ~0.2-> 2.0 Semiexposed. Seabed Overall abundance

Bay, Narrows, the Narrows); Narrows and West 8.7 = 11.2 knots,north- rapidly (>5%) slopes variable. Ampelisca

West Bank avg. = 10.1) Bank borrow pit; south into main Hudson River abdita moderately

fine-medium sand to mouth channel; West common; blue musselssilty sand elsewhere. Bank shoal and several abundant near WestLow chemical borrow pits to the west. Bank; polychaetescontamination. common in deeper,

siltier areas.

7- Ambrose 6.5-22.3 (avg. = Medium-coarse to ~21-31 5.0-11.7; avg. = 1.7-22.3; avg. ~0.5->2.0 knots, Mechanically dredged Overall abundance

Channel (~600 17.1); dredged to gravelly sand. Low 8.5 = 10.7 within channel to maintain authorized generally low, withm wide, ~7 km ~14 (45 ft) chemical depth. Slopes increase spotty settlement

long)d contamination. at sides of channel. of polychaete

Accumulations of trash Asabellides oculata.

common in the westernreach.

8- Chapel Hill 6.6-15.2 (avg. = Variable sands. Low ~20-30 5.9-11.6; avg. = 1.1-23.7; avg. ~0.5-1.3 Mechanically dredged No data, but probably

Channel (~300 10.3); dredged chemical contamination. 8.6 = 11.4 knots, east- to maintain authorized similar to Ambrose

m wide, ~7 km to ~9 (30 ft) west across depth. Slopes increase Channel.long)d channel at sides of channel. Rocks

are found in some places.

9- Raritan Bay 9.4-15.2 (avg. = Variable, silt-clay in ~20-29 5.0-12.0; avg. 0.0-24.3; avg. ~0.0-0.7 Mechanically dredged to Overall abundance

Channel (~265 13.0); dredged to west reach to mixed = 8.5 = 10.9 knots, within maintain authorized low. Ampelisca abditam wide, ~16 km ~10.5 (35 ft) mud, gravelly sand, and channel depth. Slopes increase at sometimes a dominantlong)d sand in east reach. sides of channel. taxa.

Moderate to highchemicalcontamination.

a Channel characteristics are from National Ocean Service (1995).b Sediment and benthic community descriptions are from the Cerrato et al.(1989) 1986-87 survey and Squibb et al.(1991).c Current data from National Ocean Service (1994).
Page 30Table 2a. Summary of Hudson-Raritan Estuary winter flounder diet by sampling period during 1996-97, expressed as the percent frequency of occurrence of dominant prey or

items (FO), and as the mean percent contribution of dominant prey or items to total stomach content volume (TV). (n = sample size; ES = percent of empty stomachs.)

Sampling Period

Jul 96 Oct 96 Jan 97 Apr 97 Aug 97 Nov 97

(n = 89; ES = 6.7) (n = 55; ES = 14.5) (n = 135; ES = 42.2) (n = 209; ES = 2.9) (n = 83; ES = 16.9) (n = 139; ES = 15.8)

Prey FO TV FO TV FO TV FO TV FO TV FO TV

Unidentified organic matter 24.7 5.5 34.6 11.7 5.9 20.5 14.8 2.0 55.4 21.1 30.2 6.6

Northern quahog siphons 26.9 18.5 25.5 23.9 0.7 1.3 22.5 9.0 7.2 4.9 24.5 11.0Ampelisca abdita 9.0 3.7 25.5 5.8 6.7 4.2 21.1 3.7 3.6 1.1 32.4 7.6Atlantic surfclam siphons 11.2 2.6 25.5 9.0 0.0 0.0 45.9 31.3 2.4 1.1 22.3 21.5Unidentified polychaetes 23.6 9.1 9.1 2.9 2.2 1.3 20.1 6.8 3.6 1.8 12.9 4.0Neomysis americana 1.1 1.0 3.6 2.0 41.5 23.6 3.4 0.1 1.2 0.7 2.2 0.2Unidentified clam siphons 2.2 0.9 1.8 0.3 0.7 1.3 11.5 6.9 18.1 12.7 1.4 1.7

Unciola sp. 0.0 0.0 20.0 5.8 7.4 6.8 0.5 <0.1 1.2 0.4 6.5 2.2Blue mussel juveniles 2.2 1.4 3.6 0.6 0.0 0.0 7.3 10.5 0.0 0.0 0.0 0.0Asabellides oculata 10.1 4.2 0.0 0.0 1.5 0.4 0.9 0.1 15.7 12.0 5.8 1.7Corophium sp. 10.1 3.1 10.9 5.5 0.0 0.0 3.4 0.4 1.2 0.7 1.4 <0.1Glycera sp. 14.6 11.7 1.8 1.7 0.0 0.0 3.4 1.0 2.4 6.7 2.9 2.5Unidentified hydroids 2.2 2.1 0.0 0.0 0.0 0.0 11.9 5.1 8.4 7.8 2.9 0.9Gammarus lawrencianus 6.7 4.5 5.5 2.6 5.9 6.3 1.0 0.3 0.0 0.0 3.6 1.2Atlantic rock crab juveniles 19.1 10.5 0.0 0.0 0.0 0.0 1.4 1.5 1.2 3.3 1.4 0.3Crangon septemspinosa 6.7 2.5 0.0 0.0 3.7 1.3 3.4 0.9 6.0 6.9 2.2 0.4Nemerteans 9.0 3.9 0.0 0.0 1.5 19.8 7.7 3.7 0.0 0.0 0.0 0.0
Page 31

Table 2b. Summary of Hudson-Raritan Estuary winter flounder diet by predator size group during 1996-97, expressed as the mean percent contribution of

dominant prey or items to the total stomach content volume. (n = number of nonempty stomachs included.)

Winter Flounder Size Group (cm TL)

6.0 - 9.9 10.0 - 19.9 20.0 - 29.9 30.0 - 45.0

Prey (n = 44) (n = 291) (n = 158) (n = 85)

Neomysis americana 40.81 6.20 0.01 0.00Detritus 14.47 11.80 3.40 1.70Ampelisca vadorum 8.75 5.90 3.30 0.20Unciola sp. 6.75 0.70 0.05 0.10Unidentified polychaetes 5.34 5.85 3.70 1.40Hydroids 5.34 2.32 1.90 3.50Blue mussel spat 3.69 0.63 0.46 0.65Northern quahog siphons 1.86 14.20 19.60 4.40Atlantic surfclam siphons 1.80 8.10 32.30 69.50Bivalve mollusk remains 0.85 3.01 0.74 12.50Glycera sp. 0.00 5.26 4.32 1.00Crangon septemspinosa 3.39 2.55 0.67 0.31Ensis directus 0.00 0.27 5.10 0.02
Page 32Table 2c. Summary of Hudson-Raritan Estuary juvenile winter flounder (less than 20 cm TL) diet by sampling period during 1996-97, expressed as the mean

percent contribution of dominant prey or items to total stomach content volume. (n = number of nonempty stomachs included.)

Sampling Period

Jul 96 Oct 96 Jan 97 Apr 97 Aug 97 Nov 97

Prey (n = 63) (n = 39) (n = 65) (n = 65) (n = 47) (n = 56)

Northern quahog siphons 10.50 23.77 0.01 7.68 3.59 22.15

Atlantic surfclam siphons 7.36 21.49 0.00 8.63 1.53 7.82

Ampelisca vadorum 3.98 11.20 4.28 7.03 0.93 8.62Gammarus lawrencianus 7.42 1.68 13.06 0.00 0.00 0.65

Sabellaria vulgaris 3.34 3.20 0.39 0.00 19.52 0.00

Neomysis americana 0.11 0.00 62.47 5.80 0.00 0.17Crangon septemspinosa 3.98 0.00 2.53 3.53 4.05 1.30Unciola sp. 0.00 8.00 4.46 0.06 1.29 0.07Detritus 6.64 17.68 2.96 5.14 26.10 22.15

Table 2d. Summary of Hudson-Raritan Estuary adult winter flounder (greater than or equal to 20 cm TL) diet by sampling period during 1996-97, expressed

as the mean percent contribution of dominant prey or items to total stomach content volume. (n = number of nonempty stomachs included.)

Sampling Period

Jul 96 Oct 96 Jan 97 Apr 97 Aug 97 Nov 97

Prey (n = 18) (n = 4) (n = 7) (n = 137) (n = 16) (n = 61)

Northern quahog siphons 30.73 74.77 0.00 11.71 15.41 16.82Atlantic surfclam siphons 2.74 0.00 0.00 49.95 0.00 47.79Glycera sp. 20.25 0.00 0.00 2.42 37.25 0.65Atlantic rock crab juveniles 10.93 0.00 0.00 3.55 1.28 0.11
Page 33

Table 2e. Summary of Hudson-Raritan Estuary juvenile winter flounder (6.0 - 19.9 cm TL) diet by sampling stratum during 1996-97, expressed as mean

percent contribution of dominant prey or items to total stomach content volume. (n = number of nonempty stomachs included.)

Stratum

123456789

 Prey (n = 31) (n = 46) (n = 71) (n = 51) (n = 13) (n = 70) (n = 17) (n = 18) (n = 16)Glycera sp. 15.8 9.1 5.2 0.0 0.0 2.5 0.0 0.0 0.0Sabellaria vulgaris 15.8 0.0 <1.0 <1.0 0.0 1.6 0.0 0.0 0.0Pherusa affinis 0.0 0.0 7.8 1.8 0.0 1.1 6.2 0.0 0.0Asabellides oculata 1.3 0.0 0.0 0.0 2.7 12.8 0.0 0.0 0.0Ampelisca sp. 12.0 6.2 8.3 1.4 0.0 7.0 0.0 20.8 4.9Gammarus sp. <1.0 0.0 1.4 1.2 0.0 6.8 28.0 <1.0 0.0Neomysis americana 0.0 2.2 13.3 20.5 51.1 4.1 3.5 20.0 18.8Crangon septemspinosa <1.0 0.0 4.0 5.1 9.8 2.9 0.0 0.0 0.0Cancer sp. juveniles 0.0 0.0 <1.0 15.1 0.0 1.1 2.3 0.0 0.0Northern quahog siphons 11.0 42.0 14.8 1.6 0.0 11.1 0.0 0.0 1.0Atlantic surfclam siphons 0.0 7.2 2.0 20.7 0.0 10.5 2.4 7.9 0.0Detritus 6.2 12.0 12.7 3.4 0.0 9.5 13.9 12.1 44.2

Table 2f. Summary of Hudson-Raritan Estuary adult winter flounder (20.0 - 45.0 cm TL) diet by sampling stratum during 1996-97, expressed as mean

percent contribution of dominant prey or items to total stomach content volume. (n = number of nonempty stomachs included.)

Stratum

123456789

 Prey (n = 29) (n = 30) (n = 45) (n = 39) (n = 48) (n = 18) (n = 18) (n = 5) (n = 11)Maldanopsis 20.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0Glycera sp. 12.3 11.1 3.3 0.0 0.0 <1.0 17.5 0.0 0.0Ampelisca sp. 6.9 14.5 1.0 0.0 0.0 <1.0 1.1 4.4 8.0Cancer sp. juveniles <1.0 0.0 4.0 <1.0 <1.0 20.8 1.3 0.0 0.0

Northern quahog siphons 14.5 31.6 47.1 0.0 <1.0 29.2 0.0 0.0 0.0Atlantic surfclam siphons 6.0 6.8 16.3 77.3 78.8 1.3 42.1 11.8 0.0

Ensis directus <1.0 1.1 0.0 13.4 0.0 0.0 0.0 0.0 0.0Softshell siphons 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 50.3Mulinia lateralis <1.0 1.1 <1.0 0.0 0.0 <1.0 0.0 10.0 3.3Detritus 2.0 4.9 3.5 0.0 <1.0 8.2 7.5 10.7 24.1
Page 34Table 3. Summary of other winter flounder diet studies for the coastal Middle Atlantic Bight. (Major prey taxa ranked in relative order of importance by: mean percent

frequency of occurrence (FO), mean percent contribution to total stomach content volume (TV), mean percent contribution to total stomach content weight (TW),and mean percent contribution to total stomach content dry weight (TDW), as available from each source.)

Year(s) No. Examined

(month(s), season(s), (size [TL cm]) or Primary Prey in Order of Importance

Source Study Area and/or frequency) age range, mean size) by FO, IRI, TDW, TN, TV, and/or TW

Linton (1921) Woods Hole, MA 1915-16 (May-Nov) 398 (2-22.5, ~8) TV -- amphipods, copepods, ostracods, isopods, and

“shrimp.”

Tressler and Bere Southern Long Island, 1938 (Jul-Aug) 120 (3-15) FO -- copepods, amphipods, nematodes, ostracods,(1938) NY polychaetes, and isopods.Smith (1950) Block Island Sound, RI 1948-49 (seasonally) Unknown TW -- Leptocheirus pinguis, Unciola irrorata, and Obelia

sp.Richards (1963) Long Island Sound - 1956-57 (monthly) 287 (4-17) FO? -- nemertean, Ampharete sp., L. pinguis,

sandy site Neomysis, Nereis succinea, and hydroids.

Richards (1963) Long Island Sound 1956-57 (monthly) 86 (8-16) FO? -- Nephtys incisa, L. pinguis, and Melinna cristata.

 - muddy site

de Sylva et al. (1962) Delaware Bay shore 1958 (bimonthly) 95 (3-13) FO? -- polychaetes, detritus, Edotea sp., and other isopods.

Pearcy (1962) Long Island Sound 1958-59 (monthly) 359 (1-16) TV (YOY) -- copepods to amphipods.

(Mystic River, CT) TV (ages 1+) — amphipods to polychaetes.

Mulkana (1966) Coastal Rhode Island 1962 (Jul-Oct) 124 (3-8) FO -- varies per estuary, mainly isopods (Edotea sp.),

ponds and rivers tanaids (Leptochelia sp.), polychaetes (Nereis sp. and

spionids), and amphipods (Ampelisca sp. and Lembos sp.).

Lux et al. (1996) Woods Hole, MA Sep 1961 - Dec 1962 1,248 (12-42, mostly TW -- Nereis sp., Glycera sp., Capitella sp., Macoma sp.,

(weekly) 22-38) Solemya sp., softshell siphons, Pagurus sp., Crangon, and

Ampelisca sp.

Frame (1974) Buzzards Bay, MA 1968-69 (seasonally) 176 (ages 1+) FO -- polychaetes, amphipods, and mollusks

(Nucula proxima, Tellina agilis, and Yoldia sp.).

Steimle (1985) New York Bight apex 1969-70 (summer) 196 (9-39, mostly 17-25) FO -- Pherusa affinis, Nephtys incisa, amphipods,

unidentified clam siphons, and isopods.

Festa (1979) Little Egg Harbor, NJ 1972-77 (spring-fall) 97 (3-21) TV -- Palaemonetes sp., polychaetes, clam

siphons, and nemerteans.

45 (22-33) TV -- detritus, clam siphons, polychaetes, and

Ampelisca sp.
Page 35

Kurtz (1975) Southern Long Island Oct 1973 - Jun 1974 679 (13-28) TW -- polychaetes (sabellids and terebellids), amphipods,

(NY) bays and clam siphons.

Allen et al. (1978) Hereford Inlet, NJ 1973-77 106 (4-30) FO -- polychaetes, amphipods, bivalve mollusks, Crangon,

(biweekly/monthly) and isopods.

Steimle (unpubl. data) Raritan Bay, NJ (Strata 1976 (Mar-Jun ) 84 (8-25) FO -- Asabellides oculata, spionid polychaetes, hydroids,

1-4) blue mussel spat, Glycera sp., and juvenile Ensis directus.

Homer and Boynton Chesapeake Bay, MD 1977 (May-Jun) 168 (2-15) TW -- Scolecolepides viridis, Corophium lacustra,

(1978) softshells, Macoma sp., and Nereis succinea.

Lawler, Matusky & Hudson River, NY 1979-80 (seasonally) 89 (mean = 170) TV -- polychaetes and clam siphons.

Skelly Engineers (1980)

Worobec (1984) Charles Pond, RI 1979-80 (spring-summer) 181 (11-26) TDW -- detritus, polychaetes, nematodes, and amphipods.

Conover et al. (1985) Raritan Bay, NY 1982 (monthly) 409 (15-18) TW -- A. oculata, Gammarus sp., algae, and other

amphipods.

Steimle and New York Bight apex 1982-85 (seasonally) 389 (mean = ~26) FO -- Ceriantheopsis americanus, Pherusa affinis,

Terranova (1991) Lumbrineris sp., and Nephtys sp.

Scarlett and Giust (1989) Manasquan River, NJ 1984-86 (monthly) 273 (11-40, mean = ~25) TV -- Nereis succinea, detritus, sand lance, clam

siphons, hydroids, Crangon, and polychaetes.

Scarlett (1986, 1988) Central New Jersey 1985-87 (winter-spring, 176 (12-37, means = 26-30) TV -- Nereis sp., detritus, Glycera sp., other polychaetes,

estuaries monthly) hydroids, Cyathura sp., and clam siphons.

Steimle et al. (1994) New York Bight apex 1986-89 (monthly) 3,556 (18-30) TV -- P. affinis, Ceriantheopsis americanus, Nephtys

incisa, and A. oculata.

Bharadwaj (1988) Narragansett Bay, RI 1987 (Nov) 266 (12-38) FO -- N. incisa, C. americanus, P. affinis, and Nereis sp.

Carlson (1991) New Haven Harbor, CT Sep 1989 - Nov 1990 32 (10-15) TW -- hydroids, crustaceans, and polychaetes.

65 (16-20) TW -- hydroids, snail feet, mysids, and polychaetes.

31 (21-25) TW -- Streblospio sp., hydroids, and Ampelisca sp.

19 (26-29) TW -- Crangon, Ampelisca abdita, and hydroids.

Timmons (1995) Rehoboth Bay, DE 1994 (Aug) 36 (mean = 10) TV -- polychaetes, detritus, and polychaete tubes.

Stehlik and Meise (2000) Navesink River and 1996-98 (May, Jul, and 1,399 (1.0-45.0, mean = 10) TV -- Ampelisca sp., softshell siphons, spionid and other

Sandy Hook Bay, NJ Oct) polychaetes, copepods, mysids, and Crangon.
Page 36Table 4a. Summary of Hudson-Raritan Estuary windowpane diet by sampling period during 1996-97, expressed as the percent frequency of occurrence of dominant prey or

items (FO), and as the mean percent contribution of dominant prey or items to total stomach content volume (TV). (n = sample size; ES = percent of empty stomachs.)

Sampling Period

Jul 96 Oct 96 Jan 97 Apr 97 Aug 97 Nov 97

(n = 45; ES = 2.0) (n = 59; ES = 13.5) (n = 98; ES = 33.7) (n = 163; ES = 23.3) (n = 127; ES = 11.8) (n = 78; ES = 10.3)

Prey FO TV FO TV FO TV FO TV FO TV FO TV

Neomysis americana 93.3 57.1 81.4 39.0 33.7 19.7 42.9 31.6 85.0 70.1 60.3 17.8Crangon 44.4 41.5 28.8 16.7 53.0 47.7 36.8 45.5 23.6 24.8 34.6 21.3Gammarus lawrencianus 2.2 1.1 39.0 22.8 4.1 3.0 1.2 0.1 0.8 0.5 11.5 4.8Unidentified organic matter 2.2 0.2 1.7 <0.1 0.0 0.0 15.3 1.9 2.4 1.6 5.1 1.1Ampelisca abdita 0.0 0.0 3.4 2.9 1.0 0.9 3.7 1.6 0.8 0.1 9.0 7.1Pseudodiaptomus coronatus 0.0 0.0 8.5 7.7 0.0 0.0 0.6 1.4 0.0 0.0 2.6 1.9Unidentified juvenile fish 0.0 0.0 3.4 1.6 1.0 <0.1 1.2 2.5 0.0 0.0 10.3 3.7Juvenile Anchoa sp. 0.0 0.0 0.0 0.0 0.0 0.0 2.5 5.0 0.0 0.0 12.8 7.9
Page 37

Table 4b. Summary of Hudson-Raritan Estuary windowpane diet by predator size group during 1996-97, expressed as the percent frequency of occurrence of dominant prey or items (FO), and as the mean percent contribution of dominant prey or items to total stomach content volume (TV).

(n = number of nonempty stomachs included.)

Windowpane Size Group (cm TL)

<9.9 10.0 - 19.9 20.0 - 29.9 >30.0

(n = 126) (n = 178) (n = 229) (n = 24)

Prey FO TV FO TV FO TV FO TV

Neomysis americana 100.0 91.1 88.2 61.6 65.9 38.5 58.3 23.2

Crangon septemspinosa 7.7 4.9 35.4 23.6 54.6 51.0 66.7 56.3Gammarus sp. 0.0 0.0 11.8 1.5 7.4 2.4 8.3 1.6Anchoa sp. 0.0 0.0 5.6 12.3 1.3 1.4 4.2 8.7Detritus 7.7 1.8 1.7 <1.0 12.7 <1.0 0.0 0.0
Page 38Table 4c. Summary of Hudson-Raritan Estuary windowpane diet by sampling stratum during 1996-97, expressed as mean percent contribution of

dominant prey or items to total stomach content volume. (n = number of nonempty stomachs included.)

Stratum

123456789

Prey (n = 23) (n = 24) (n = 34) (n = 18) (n = 33) (n = 25) (n = 24) (n = 16) (n = 32)

Neomysis 49.5 17.3 35.7 30.8 14.3 72.2 82.6 32.5 23.4Crangon 44.4 77.7 52.8 21.2 36.2 26.1 6.1 64.6 76.5

Table 5. Summary of other windowpane diet studies for the coastal Middle Atlantic Bight. (Major prey taxa ranked in relative order of importance by: mean percent frequency

of occurrence (FO), mean percent contribution to total stomach content volume (TV), and mean percent contribution to total stomach content weight (TW), asavailable from each source.)

Year(s) No. Examined

(month(s), season(s), (size [TL cm]) or Primary Prey in Order of Importance

Source Study Area and/or frequency) age range, mean size) by FO, IRI, TDW, TN, TV, and/or TW

Moore (1947) Long Island - Block 1943-44 (monthly) 654 (18-34) TV -- Neomysis, Crangon, chaetognaths, and larval fish

Island Sounds (sand lance and silver hake).

Smith (1950) Block Island Sound 1948-49 (seasonally) Unknown, ~200 TW -- Neomysis, small fish, and squid.

Richards (1963) Long Island Sound 1955-56 (seasonally) Sand station - 49 (3-13) FO -- Neomysis and Crangon.

Mud station - 25 (3-13) FO -- Neomysis.

de Sylva et al. (1962) Delaware Bay shore 1958-60 (Feb-Jul) 31 (5-17) FO -- Neomysis, Crangon, and copepods.Langton and Bowman New Jersey - North 1969-72 (spring/fall) 163 (mean = 25) TW -- Neomysis, pandalid shrimp, Crangon, and other

(1981) Carolina decapod crustaceans.Kimmel (1973) Chesapeake Bay 1971-72 (spring) 16 (7-17) TV -- Neomysis, larval bay anchovy, and Crangon.

mouth 18 (19-24) TV -- Neomysis, Crangon, and bay anchovy.

Hickey (1975) Eastern Long 1972 (Apr-Dec) 120 (6-26) TV -- Neomysis, Crangon, fish eggs, larvae, and “bait.”

Island Sound
Page 39

Festa (1979) Little Egg Harbor, NJ 1972-77 (summer/fall) 4 (15-34) TV -- Crangon, sand lance, Neomysis, and detritus.

FO -- Detritus, Neomysis, and Crangon.

Allen et al. (1978) Hereford Inlet, NJ 1973-77 (monthly) 60 (6-27) FO -- Mysids, Crangon, amphipods, and decapod crab larvae.

Steimle and Terranova (1991) New York Bight apex 1982-85 (seasonally) 131 (mean = ~27) FO -- Mysids (Neomysis mostly), Crangon, and

Dichelopandalus leptocerus.

Warkentine and Rachlin (1988) New Jersey coast Unreported Unreported FO -- Neomysis.

Carlson (1991) New Haven Harbor, Sep 1989 - Nov 1990 96 (11-34) TW -- Crangon, bay anchovy, and naked goby (Gobiosoma

CT (monthly) bosc).

FO -- Crangon, bay anchovy, goby, and Neomysis.
Page 40Table 6. Summary of Hudson-Raritan Estuary little skate diet by sampling period during 1996-97, expressed as the percent frequency of occurrence of dominant prey or items

(FO), and as the mean percent contribution of dominant prey or items to total stomach content volume (TV). (n = sample size; ES = percent of empty stomachs.)

Sampling Period

Jul 96 Oct 96 Jan 97 Apr 97 Aug 97 Nov 97

(n = 10; ES = 0) (n = 33; ES = 0) (n = 85; ES = 0) (n = 116; ES = 0.8) (n = 0; ES = N/A) (n = 88; ES = 1.1)

Prey FO TV FO TV FO TV FO TV FO TV FO TV

Crangon septemspinosa 92.9 90.1 81.7 29.9 85.9 36.2 87.9 31.2 - - 77.2 20.3Juvenile Atlantic rock crab 7.1 4.7 24.2 9.0 75.3 38.1 44.0 19.9 - - 35.2 9.7Juvenile Ovalipes ocellatus 7.1 1.4 36.4 15.6 1.2 0.2 1.7 0.7 - - 25.0 6.4Unidentified organic matter 14.3 2.6 24.2 7.3 2.4 0.5 3.4 1.0 - - 6.8 1.1Ensis directus 0.0 0.0 3.0 1.8 2.4 0.6 17.2 6.1 - - 18.2 3.8Neomysis americana 7.1 1.2 6.1 0.6 15.3 2.4 28.5 9.3 - - 13.6 2.8Unidentified juvenile flounder 0.0 0.0 6.1 1.9 3.5 0.7 0.9 0.6 - - 13.6 5.4Juvenile blue crab 0.0 0.0 0.0 0.0 0.0 0.0 4.3 1.4 - - 28.4 8.8Pagurus sp. 0.0 0.0 6.1 0.7 1.2 0.4 5.2 1.6 - - 18.1 5.9Atlantic surfclam 0.0 0.0 6.1 9.5 0.0 0.0 4.3 1.6 - - 3.4 0.5Gammarus lawrencianus 0.0 0.0 6.1 6.0 2.4 0.4 0.0 0.0 - - 6.8 1.1

Table 7. Summary of other little skate diet studies for the coastal Middle Atlantic Bight. (Major prey taxa ranked in relative order of importance by: mean percent frequency

of occurrence (FO), mean percent contribution to total stomach content volume (TV), mean percent contribution to total stomach content weight (TW), meanpercent contribution to total number of individual items in the stomach (TN), and an index of relative importance (IRI), as available from each source.)

Year(s) No. Examined

(month(s), season(s), (size [TL cm]) or Primary Prey in Order of Importance

Source Study Area and/or frequency) age range, mean size) by FO, IRI, TDW, TN, TV, and/or TW

Linton (1901) Woods Hole, MA 1899 (summer) 32 (unknown) FO? -- “Hermit, rock, blue, and mud crabs, shrimp,

amphipods, polychaetes, squid, clams, fish.”

Field (1907) Woods Hole, MA 1905 (summer) 516 (~30-60) FO -- Crabs (rock, lady, mud, and hermit), Crangon, fish

(sand lance, butterfish, herring, and bothid flounder),squid, Nereis sp., Ensis directus, and amphipods.
Page 41

Smith (1950) Block Island Sound 1948-49 (seasonally) Unknown, ~200 TV -- Leptocheirus pinguis, Atlantic rock crab, Crangon,

(unknown) other crustaceans, polychaetes, mollusks, small fish, and

squid.

Fitz and Daiber Delaware Bay 1954-55 (seasonally) 185 (mean = 45) FO -- Crangon, Lepidonotus squamatus, E. directus,

(1963) Nereis sp., Pagurus sp., and fish (northern searobin and

windowpane).

TN -- Crangon and E. directus.

Richards et al. Long Island Sound 1956-57 (seasonally) 3 (9-11) FO -- Copepods, Neomysis, Heteromysis formosa, and

(1963) Crangon.

Vinogradov (1984) Georges Bank - 1968-74 (unreported) 253 (unreported) TW -- Crustaceans, polychaetes, and fish.

Southern New England

McEachlan et al. (1976) Offshore 1969-70 (seasonally) 785 (mean = ~45-50) IRI -- Decapod crustaceans (Crangon, etc.), amphipods

(Leptocheirus pinguis, etc.), and polychaetes (Eunice sp.

and Nereis sp.).

Bowman et al. (1987) Southern New England 1973-76 (spring/fall) 168 (16-20) TW -- Crangon, Atlantic rock crab, and other crustaceans.

Sedberry (1983) New York Bight 1976-77 (seasonally) 1,050 (5-30, disc width) TV -- Atlantic rock crab, Unciola irroratus, Byblis

serrata,

and juvenile red hake.

IRI -- Amphipods and decapod crustaceans.

Carlson (1991) New Haven Harbor, CT Sep 1989 - Nov 1990 115 (31-57) TW -- Atlantic rock crab, decapod crustacean fragments,

Ovalipes ocellatus, fish, and Crangon.

FO -- Atlantic rock crab, Crangon, O. ocellatus, and

Pagurus sp.
Page 42Table 8. Summary of Hudson-Raritan Estuary scup diet by sampling period during 1996-97, expressed as the percent frequency of occurrence of dominant prey or items

(FO), and as the mean percent contribution of dominant prey or items to total stomach content volume (TV). (n = sample size; ES = percent of empty stomachs.)

Sampling Period

Jul 96 Oct 96 Jan 97 Apr 97 Aug 97 Nov 97

(n = 58; ES = 7.0) (n = 45; ES = 4.0) (n = 0; ES = N/A) (n = 76; ES = 1.0) (n = 75; ES = 1.0) (n = 0; ES = N/A)

Prey FO TV FO TV FO TV FO TV FO TV FO TV

Unidentified organic matter 36.2 14.7 37.8 8.9 - - 46.1 14.8 21.3 3.4 - -Unidentified bivalve remains 22.4 11.2 8.9 7.3 - - 1.3 0.4 26.7 8.0 - -Gammarus lawrencianus 48.3 20.8 17.8 10.4 - - 0.0 0.0 0.0 0.0 - -Neomysis americana 50.0 24.4 42.2 13.5 - - 19.8 9.2 17.3 2.8 - -Crangon septemspinosa 8.6 6.1 17.8 4.6 - - 3.9 2.1 30.7 15.5 - -Ampelisca abdita 0.0 0.0 13.3 3.7 - - 9.2 4.6 18.0 4.6 - -Unidentified polychaetes 6.9 4.1 6.7 2.8 - - 35.5 15.2 9.3 2.2 - -Juvenile Atlantic rock crab 12.1 5.1 2.2 0.9 - - 2.6 1.1 21.3 9.2 - -Juvenile blue mussel 1.7 0.5 2.2 0.3 - - 31.6 10.2 1.3 0.2 - -Northern quahog siphons 1.7 1.5 13.3 7.0 - - 0.0 0.0 17.3 9.8 - -Pagurus spp. 12.1 8.4 15.6 5.8 - - 2.6 2.1 13.3 5.0 - -Ensis directus 1.7 1.0 0.0 0.0 - - 1.3 7.4 13.3 8.8 - -

Table 9. Summary of other scup diet studies for the coastal Middle Atlantic Bight. (Major prey taxa ranked in relative order of importance by: mean percent frequency of

occurrence (FO), mean percent contribution to total stomach content volume (TV), mean percent contribution to total stomach content weight (TW),mean percentcontribution to total stomach content dry weight (TDW), and an index of relative importance (IRI), as available from each source.)

Year(s) No. Examined

(month(s), season(s), (size [TL cm]) or Primary Prey in Order of Importance

Source Study Area and/or frequency) age range, mean size) by FO, IRI, TDW, TN, TV, and/or TW

Linton (1901) Woods Hole, MA 1896-99 (summer) 51 (“YOY”) FO? -- Copepods and small crustaceans.

58 (“adults”) FO? -- Small fish, squid, polychaetes, crabs, shrimp,

amphipods, mollusks, and hydroids.

Richards (1963) Long Island Sound - 1956-57 (seasonally) 167 (2-15) FO -- Polychaetes (Ampharete sp.), copepods

sand station (Pseudodiaptomus coronatus), and amphipods (Photis

sp. and Leptocheirus pinguis).
Page 43

Bowman et al. (1987) Southern New England 1973-76 (spring/fall) 367 (<15) TW -- Polychaetes, amphipods, decapod crustaceans

(Atlantic rock crab), and squid.

Sedberry (1983) New York Bight 1976-77 (seasonally) 138 (5-15) IRI -- Amphipods (hyperids and gammarids) and

polychaetes (Potamilla reniforma and Glycera)

102 (15-30) IRI -- Polychaetes, gammarid amphipods (Ericthonius sp.),

and decapod crustaceans (Atlantic rock crab)

Steimle (unpubl. data) Raritan Bay 1976 (Mar-Jun) 13 (YOY) FO -- Asabellides oculata, copepods, polydorid

polychaetes, Mulinia lateralis, blue mussel spat, and

hydroids.

Michelman (1988) Narragansett Bay, RI 1987 (seasonally) 66 (“juveniles”) TDW -- Polychaetes (maldanids, Nephtys incisa, Nereis

sp., and Pherusa affinis), Pagurus sp., Neomysis,

amphipods (L. pinguis, etc.), mollusks, Ceriantheopsisamericanus, and fish larvae.

Steimle et al. Delaware Bay 1989-94 (May/Jun, 246 (9.0-28.0, mean = TV -- blue mussels, Ensis directus, unidentified mollusks,

(unpubl. data) Aug/Sep) 14.2) Metridium senile, Dyspanopeus sayi, Neomysis, Atlantic

rock crab, and unidentified fish remains.
Page 44Table 10. Summary of Hudson-Raritan Estuary summer flounder diet by sampling period during 1996-97, expressed as the percent frequency of occurrence of dominant prey

or items (FO), and as the mean percent contribution of dominant prey or items to total stomach content volume (TV). (n = sample size; ES = percent of emptystomachs.)

Sampling Period

Jul 96 Oct 96 Jan 97 Apr 97 Aug 97 Nov 97

(n = 50; ES = 10.0) (n = 8; ES = 25.0) (n = 5; ES = 40.0) (n = 38; ES = 50.0) (n = 128; ES = 25.0) (n = 0; ES = N/A)

Prey FO TV FO TV FO TV FO TV FO TV FO TV

Crangon septemspinosa 78.0 44.0 75.0 41.2 60.0 89.3 36.8 31.4 34.4 12.2 - -

Neomysis americana 30.0 19.6 0.0 0.0 20.0 10.7 5.3 3.1 33.6 11.5 - -Unidentified juvenile fish 14.0 9.1 12.5 21.0 0.0 0.0 13.2 10.9 12.5 4.3 - -Juvenile Ovalipes ocellatus 2.0 1.2 0.0 0.0 0.0 0.0 0.0 0.0 32.0 17.1 - -Unidentified organic matter 2.0 0.3 0.0 0.0 0.0 0.0 5.3 0.2 8.6 3.9 - -Unidentified juvenile flounder 0.0 0.0 0.0 0.0 0.0 0.0 7.9 24.0 2.3 1.7 - -Juvenile windowpane 4.0 9.5 0.0 0.0 0.0 0.0 0.0 0.0 1.5 6.5 - -Juvenile Atlantic rock crab 4.0 7.2 0.0 0.0 0.0 0.0 0.0 0.0 7.8 7.0 - -

Table 11. Summary of other summer flounder diet studies for the coastal Middle Atlantic Bight. (Major prey taxa ranked in relative order of importance by: mean percent

frequency of occurrence (FO), mean percent contribution to total stomach content volume (TV), and mean percent contribution to total stomach content weight(TW), as available from each source.)

Year(s) No. Examined

(month(s), season(s), (size [TL cm]) or Primary Prey in Order of Importance

Source Study Area and/or frequency) age range, mean size) by FO, IRI, TDW, TN, TV, and/or TW

Linton (1901) Woods Hole, MA Unknown 24 (unknown) FO -- Squid and fish (e.g., juvenile scup).

Poole (1964) Great South Bay, NY 1958-59 (summer/fall) 1,210 (25-67, mostly TW -- Crangon, juvenile winter flounder, juvenile blue

25-42) crabs, Neomysis, and northern pipefish.

FO -- Crangon, Neomysis, and northern pipefish.

de Sylva et al. (1962) Delaware Bay shore 1958 (bimonthly) 26 (5-21) FO -- Neomysis, Crangon, Menidia, unidentified fish, and

other crustaceans.

Langton and New Jersey - North 1969-72 (spring/fall) 44 (mean = 38) TW -- Loligo squid, juvenile silver hake and scup, and

Bowman (1981) Carolina Atlantic rock crab.
Page 45

Kimmel (1973) Mouth of Chesapeake Bay 1971-72 (spring/summer) 35 (4-20) TV -- Neomysis and goby.

36 (20-48) TV -- Neomysis, Crangon, and bay anchovy.

Festa (1979) Little Egg Harbor, NJ 1972-77 (summer/fall) 25 (6-24) TV -- Fish (anchovy), Crangon, Palaemonetes sp., and

Neomysis.

13 (26-65) TV -- Fish (juvenile searobin and winter flounder, silver

perch, anchovy), and juvenile blue crab.

Smith and Daiber Delaware Bay Unknown 131 (31-73) TV -- Juvenile weakfish, Crangon, Neomysis, bay

(1977) anchovy, squid, Menidia, herring, and Pagurus sp.

Allen et al. (1978) Hereford Inlet, NJ 1973-77 57 (1-55) FO -- Crangon, mysids, and fish.

(biweekly/monthly)

Homer and Boynton Chesapeake Bay, MD 1977 (summer) 198 (5-27) TW -- Juvenile weakfish and spot, bay anchovy, juvenile

(1978) Atlantic menhaden, Crangon, amphipods, juvenile

croaker, mollusks, Menidia, Neomysis, and nematodes.

Steimle and New York Bight apex 1982-85 (seasonally) 20 (mean = ~30) FO -- Neomysis, fish, and amphipods.

Terranova (1991)Scarlett and Giust Manasquan River, NJ 1984-86 (monthly) 90 (19-44, mean = ~31) TV -- Juvenile blue crab, Crangon, and fish (mummichog,(1989) sand lance, juvenile winter flounder, and unidentified species).Rountree and Able Little Egg Harbor, NJ 1987-90 (summer) 137 (17-31, mean = 23) TW -- Menidia, mummichog, Palaemonetes sp., and

(1992) Crangon.

FO -- Crangon, Menidia, Palaemonetes sp., andmummichog.

Steimle et al. Delaware Bay 1989-94 (May/Jun and 43 (23.0-56.0, mean = TV -- Neomysis and unidentified fish and mollusk

(unpubl. data) Aug/Sep) 35.3) remains.Timmons (1995) Rehoboth Bay, DE 1994 (summer) 36 (8-25, mean = 13) TV -- Palaemonetes sp., juvenile blue crab, other crabs

and shrimp, and mysids.
Page 46Table 12. Summary of Hudson-Raritan Estuary red hake diet by sampling period during 1996-97, expressed as the percent frequency of occurrence of dominant prey or items

(FO), and as the mean percent contribution of dominant prey or items to total stomach content volume (TV). (n = sample size; ES = percent of empty stomachs.)

Sampling Period

Jul 96 Oct 96 Jan 97 Apr 97 Aug 97 Nov 97

(n = 1; ES = 0) (n = 0; ES = N/A) (n = 62; ES = 5) (n = 64; ES = 0) (n = 7; ES = 14) (n = 32; ES = 0)

Prey FO TV FO TV FO TV FO TV FO TV FO TV

Crangon septemspinosa 100.0 50.0 - - 85.5 47.4 91.2 46.2 71.4 47.5 56.3 23.3

Gammarus lawrencianus 100.0 50.0 - - 8.1 9.5 10.3 1.5 0.0 0.0 43.8 18.8Neomysis americana 0.0 0.0 - - 48.4 30.3 39.7 14.5 28.6 26.2 12.5 2.4Unidentified organic matter 0.0 0.0 - - 8.1 0.9 20.6 6.2 14.3 3.3 3.1 1.0Juvenile Atlantic rock crab 0.0 0.0 - - 1.6 1.0 7.3 1.1 0.0 0.0 6.3 2.8Unidentified fish 0.0 0.0 - - 0.0 0.0 1.5 3.3 0.0 0.0 12.5 3.7Dichelopandalus leptocerus 0.0 0.0 - - 0.0 0.0 0.0 0.0 14.3 23.0 0.0 0.0

Table 13. Summary of other red hake diet studies for the coastal Middle Atlantic Bight. (Major prey taxa ranked in relative order of importance by: mean percent frequency

of occurrence (FO), mean percent contribution to total stomach content volume (TV), mean percent contribution to total stomach content weight (TW), meanpercent contribution to total number of individual items in the stomach (TN), and an index of relative importance (IRI), as available from each source.)

Year(s) No. Examined

(month(s), season(s), (size [TL cm]) or Primary Prey in Order of Importance

Source Study Area and/or frequency) age range, mean size) by FO, IRI, TDW, TN, TV, and/or TW

Richards (1963) Long Island Sound 1956-57 (seasonally) Sand station - 21 (9-20) FO -- Polychaetes (Glycera sp.), amphipods (Ampelisca sp.

and Leptocheirus pinguis), Neomysis, Heteromysis formosa,and Crangon.

Steimle and Ogren New York Bight 1966-67 (Jul-Aug) 31 (mostly 33-36) TV -- Crangon, unidentified tunicate, Atlantic rock crab,(1982) unidentified fish, and Nereis sp.

Steimle (1985) New York Bight 1968-70 (summer) 219 (mostly 17-25) FO -- Crangon, polychaetes (Pherusa affinis and Nephtys

incisa), Neomysis, and amphipods (Unciola sp. and L.pinguis).

Vinogradov (1984) Southern New England 1968-74 (spring/fall?) 1,892 (unreported) TW -- Sipunculids, unidentified fish, crabs, squid, “shrimp,”

gammaridean amphipods, and bivalve mollusks.
Page 47

Bowman et al. (1987) Southern New England 1973-76 (spring/ fall) “208” (<10) TW -- Crustaceans (Dichelopandalus leptocerus),

chaetognaths, amphipods, and decapod crustaceans.

Part of above (10-20) TW -- Crustaceans, amphipods (Gammarus sp.), decapod

crustaceans (Crangon), and euphausids.

Steimle (unpubl. data) Raritan Bay 1976 (Mar-Jun) 45 (Subadult) FO -- Crangon.

Sedberry (1983) New York Bight 1976-77 (seasonally) 716 (5-20) IRI -- Amphipods (Unciola irrorata) and copepods.

425 (25-50) IRI -- Amphipods (U. irrorata), Atlantic rock crab, fish, and

ocean scallops.

Luczkovich and Coastal New Jersey 1979-80 (1 yr, monthly) 130 (2-9) FO -- Unidentified crustaceans, decapod crustaceansOlla (1983) (Crangon), amphipods (Unciola sp., L. pinguis,

Monoculodes sp., and Ericthonius sp.), calanoid copepods,and mysids.

Steimle and New York Bight apex 1982-85 (seasonally) 144 (not noted) FO -- Crangon, P. affinis, juvenile Atlantic rock crab, sand

Terranova (1991) lance (Ammodytes sp.), and D. leptocerus.

Steimle (1994) New York Bight apex 1986-89 (monthly) 1,047 (10-34, FO -- P. affinis, Crangon, D. leptocerus, Atlantic rock crab,

mostly >17) and N. incisa.

Rachlin and Central New Jersey Unreported 133 (6-32, mostly TN -- Neomysis, decapod crustaceans (Crangon),

Warkentine (1988) subadults) nematodes, copepods, amphipods, isopods, and fish.
Page 48Table 14. Summary of Hudson-Raritan Estuary weakfish diet by sampling period during 1996-97, expressed as the percent frequency of occurrence of dominant prey or items

(FO), and as the mean percent contribution of dominant prey or items to total stomach content volume (TV). (n = sample size; ES = percent of empty stomachs.)

Sampling Period

Jul 96 Oct 96 Jan 97 Apr 97 Aug 97 Nov 97

(n = 0; ES = N/A) (n = 66; ES = 12) (n = 0; ES = N/A) (n = 0; ES = N/A) (n = 117; ES = 3) (n = 14; ES = 57)

Prey FO TV FO TV FO TV FO TV FO TV FO TV

Crangon septemspinosa - - 63.6 20.5 ---- 54.7 30.0 14.3 23.9Neomysis americana - - 31.8 3.3 ---- 81.3 26.7 14.3 4.5Unidentified fish (Anchoa?) - - 24.2 3.8 ---- 21.4 12.5 7.1 15.9Gammarus lawrencianus - - 27.3 4.3 - - - - 0.9 0.9 14.3 11.4Unidentified organic matter - - 10.6 0.5 - - - - 1.7 0.9 7.1 1.1Anchoa mitchilli - - 13.5 17.3 - - - - 1.9 <0.1 0.0 0.0Ampelisca abdita - - 0.0 0.0 - - - - 1.7 <0.1 7.1 3.4

Juvenile silver hake - - 0.0 0.0 - - - - 0.0 0.0 7.1 36.4

Table 15. Summary of other weakfish diet studies for the coastal Middle Atlantic Bight. (Major prey taxa ranked in relative order of importance by: mean percent frequency of

occurrence (FO), mean percent contribution to total stomach content volume (TV), and mean percent contribution to total stomach content weight (TW), as availablefrom each source.)

Year(s) No. Examined

(month(s), season(s), (size [TL cm]) or Primary Prey in Order of Importance

Source Study Area and/or frequency) age range, mean size) by FO, IRI, TDW, TN, TV, and/or TW

Welsh and Breder (1924) Acushnet River, MA 1882 (Sep) 28 (7-11) TV -- Fish (killifish and river herring) and “shrimp.”

Peck (1896) Woods Hole, MA 1893 (summer) 570 (“older fish”) FO -- Juvenile herring and bluefish, butterfish, herring, squid,

crustaceans, and juvenile Atlantic menhaden.

Welsh and Breder (1924) Cape May, NJ 1916 (Aug) 32 (3-8) TV -- Mysids, “shrimp,” unidentified crustaceans, and juvenile

herring.

1919 (May) 30 (24-39) TV -- Mysids, unidentified crustaceans, and “shrimp.”

Welsh and Breder (1924) Cape Charles, VA 1916 (Sep) 45 (4-12) TV -- Mysids, copepods, amphipods, and juvenile herring.

Breder (1922b) Raritan Bay, NJ 1921 (summer) ? (33-51) FO? -- Squid, shrimp, juvenile Atlantic menhaden, silver perch,

and anchovy.

Tressler and Bere (1939) Southern Long 1938 (summer) 89 (4-11) FO? -- Crangon and copepods.

Island, NY
Page 49

Shuster (1959) Delaware Bay 1952 (summer/fall) 205 (12-30) FO -- Neomysis, unidentified fish, Solen viridis, and Crangon

Richards (1963) Long Island Sound 1956-57 (seasonally) Sand station - 23 (4-10) FO? -- Crangon, Upogebia sp., juvenile bay anchovy,

copepods, and Ampelisca sp.

Mud station - 48 (4-8) FO? -- Crangon, Neomysis, and copepods.

de Sylva et al. (1962) Delaware Bay 1958-60 (summer/fall) 220 (2-15) FO -- Neomysis, Crangon, decapod larvae, juvenile Limulus

polyphemus, and unidentified juvenile fish.

Van Engle and Chesapeake Bay, VA 1967 (summer) 268 (“juveniles”) TV -- Unidentified fish (mostly bay anchovy and naked goby),Joseph (1968) Neomysis, Crangon, copepods, and amphipods.

FO -- Neomysis, fish, copepods, and amphipods.

Thomas (1971) Delaware River 1969 (summer/fall) 494 (5-18) FO -- Neomysis, Gammarus sp., unidentified fish , juvenile

weakfish, detritus, and copepods.

64 (>18) FO -- Juvenile weakfish, unidentified fish, Neomysis, and

Gammarus sp.

Kimmel (1973) Mouth of Chesapeake 1971-72 (spring/summer) 27 (2-28) TV -- Neomysis, calanoid copepods, Crangon, and unidentified

Bay crustaceans.

Festa (1979) Little Egg Harbor, NJ 1972-77 (summer/fall) 51 (<17) TV -- Neomysis, fish (anchovy), Crangon, and Ampelisca sp.

FO -- Neomysis, Crangon, fish, and Ampelisca sp.

Chao and Musick (1977) York River, VA 1973 (summer) 36 (7-18) FO -- Bay anchovy, Neomysis, and other fish.

Allen et al. (1978) Hereford Inlet, NJ 1973-77 (biweekly/ 86 (3-22) FO -- Mysids, Crangon, crab larvae, and unidentified fish.

monthly)

Bason et al. (1975, Chesapeake and 1974-76 (summer/fall) 1,119 (2-44, TV -- Neomysis, bay anchovy, Crangon, Gammarus sp.,

1976); Keirsey et al. Delaware Canal, DE mostly < 10) juvenile fish (Atlantic menhaden, weakfish, and croaker),

(1977) polychaetes, and Corophium sp.

Homer and Boynton Chesapeake Bay, MD 1977 (summer) 808 (3-19) TW -- Bay anchovy, Pseudodiaptomus coronatus, juvenile

(1978) weakfish, Nereis succinea, and other polychaetes.Scarlett and Giust Manasquan River, NJ 1984-86 (monthly) 45 (9-20) TV -- Crangon, sand lance, detritus, Neomysis, unidentified(1989) fish, and naked goby.Hartman and Brandt Chesapeake Bay 1990-92 (bimonthly?) 564 (age 0) TW -- Juvenile bay anchovy and other fish (e.g. juvenile(1995) croaker).

353 (age 1) TW -- Bay anchovy, juvenile Atlantic menhaden, spot, and

other fish.

 54 (age 2+) TW -- Spot, Atlantic menhaden, juvenile summer flounder,

and bay anchovy.
Page 50Table 16. Summary of Hudson-Raritan Estuary spotted hake diet by sampling period during 1996-97, expressed as the percent frequency of occurrence of dominant prey or

items (FO), and as the mean percent contribution of dominant prey or items to total stomach content volume (TV). (n = sample size; ES = percent of empty stomachs.)

Sampling Period

Jul 96 Oct 96 Jan 97 Apr 97 Aug 97 Nov 97

(n = 44; ES = 0) (n = 14; ES = 7) (n = 1; ES = 0) (n = 72; ES = 0) (n = 3; ES = 33) (n = 28; ES = 0)

Prey FO TV FO TV FO TV FO TV FO TV FO TV

Crangon septemspinosa 95.5 60.9 85.7 42.4 100.0 33.3 76.4 47.1 66.7 38.9 89.3 31.3Neomysis americana 15.9 5.9 14.3 4.1 100.0 33.3 33.3 8.9 100.0 40.7 7.1 0.3Unidentified juvenile fish 0.0 0.0 7.1 13.5 0.0 0.0 5.6 5.0 33.3 20.4 50.0 17.2Unidentified organic matter 6.8 0.4 0.0 0.0 0.0 0.0 13.9 5.4 0.0 0.0 3.6 <0.1Gammarus lawrencianus 15.9 4.4 28.6 9.3 0.0 0.0 0.0 0.0 0.0 0.0 17.9 4.3Pseudodiaptomus sp. 0.0 0.0 0.0 0.0 100.0 33.3 13.9 2.1 0.0 0.0 10.7 0.1Unidentified juvenile flounder 2.3 0.5 0.0 0.0 0.0 0.0 1.4 4.1 0.0 0.0 17.9 4.2Juvenile Anguilla rostrata 0.0 0.0 7.1 13.5 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0Unidentified hydroids 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 17.9 9.5

Table 17. Summary of other spotted hake diet studies for the coastal Middle Atlantic Bight. (Major prey taxa ranked in relative order of importance by: mean percent frequency

of occurrence (FO), mean percent contribution to total stomach content volume (TV), mean percent contribution to total stomach content weight (TW), and meanpercent contribution to total number of individual items in the stomach (TN), as available from each source.)

Year(s) No. Examined

(month(s), season(s), (size [TL cm]) or Primary Prey in Order of Importance

Source Study Area and/or frequency) age range, mean size) by FO, IRI, TDW, TN, TV, and/or TW

Linton, quoted in Woods Hole, MA? Unknown 141 (unknown) FO? -- “Mysis,” shrimp, crabs, amphipods, isopods, fish,

Hildebrand and polychaetes, leeches, sponges, and hydroids.Schroeder (1928)Richards (1963) Long Island Sound 1955-57 (monthly) Sand station - 17 (6-20) FO -- Leptocheirus pinguis, Crangon, and sand lance.

Mud station - 16 (5-19) FO -- Sand lance, Corophium sp., and copepods (Temora

sp.).

Barans (1969) Chesapeake Bay 1966-68 (Mar-Jul) 600 (5-18) FO -- Crangon, other shrimp, Neomysis, squid, polychaetes,

and unidentified fish.

Kimmel (1973) Mouth of Chesapeake 1971-72 (spring) 44 (3-20) TV -- Neomysis, Crangon, Ampelisca sp., other crustaceans,

Bay and bay anchovy.
Page 51

Festa (1979) Little Egg Harbor, NJ 1972-77 (spring/fall) 16 (5-19) TV -- Crangon, Palaemonetes sp., and Neomysis.

FO -- Crangon and Neomysis.

Bowman et al. (1987) Southern New England 1973-76 (spring/fall) 244 (YOY) TW -- Euphausids, Crangon, and Dichelopandalus

leptocerus.

Rachlin and Coastal New York Bight 1973-74 (summer) 156 (5-21) TN -- Neomysis, copepods (Acartia sp. and Temora sp.),

Warkentine (1987) Crangon, amphipods (Gammarus sp.), round herring

(Etrumeus teres), and decapod crustacean larvae.

Allen et al. (1978) Hereford Inlet, NJ 1973-77 (biweekly/ 101 (5-17) FO -- Crangon, mysids, detritus, and copepods.

monthly)

Sedberry (1983) New York Bight shelf 1976-77 (seasonally) 129 (< 20) TV -- Atlantic rock crab, D. leptocerus, Crangon, unidentified

hake, Gulf Stream flounder (Citharichthys arctifrons), and

amphipods.

68 (>20) TV -- As above, plus squid and juvenile sea scallops.

Homer and Boynton Chesapeake Bay, MD 1977 (spring) 19 (10-15) TW -- Crangon, juvenile Atlantic menhaden, Nereis

(1978) succinea, bay anchovy, and juvenile spot.
Page 52Table 18. Summary of Hudson-Raritan Estuary striped searobin diet by sampling period during 1996-97, expressed as the percent frequency of occurrence of dominant prey

or items (FO), and as the mean percent contribution of dominant prey or items to total stomach content volume (TV). (n = sample size; ES = percent of emptystomachs.)

Sampling Period

Jul 96 Oct 96 Jan 97 Apr 97 Aug 97 Nov 97

(n = 26; ES = 4) (n = 39; ES = 11) (n = 0; ES = N/A) (n = 5; ES = 40) (n = 81; ES = 5) (n = 2; ES = 0)

Prey FO TV FO TV FO TV FO TV FO TV FO TV

Crangon septemspinosa 96.2 56.2 74.4 42.3 - - 60.0 71.1 75.3 37.4 50.0 71.4Neomysis americana 23.1 12.8 25.6 9.5 - - 0.0 0.0 37.0 10.2 0.0 0.0Unidentified organic matter 0.0 0.0 15.4 8.6 - - 0.0 0.0 6.2 0.7 0.0 0.0Juvenile Atlantic rock crab 3.8 2.7 7.7 9.0 - - 0.0 0.0 11.1 9.7 0.0 0.0Coal pebbles 3.8 0.7 2.6 <0.1 - - 0.0 0.0 4.9 1.6 0.0 0.0Juvenile Ovalipes ocellatus 0.0 0.0 5.1 8.1 - - 0.0 0.0 9.9 5.4 0.0 0.0Unidentified juvenile fish 3.8 7.7 2.6 0.9 - - 0.0 0.0 11.1 5.8 0.0 0.0Juvenile striped searobin 0.0 0.0 0.0 0.0 - - 0.0 0.0 7.4 8.8 0.0 0.0Ampelisca abdita 7.7 1.3 0.0 0.0 - - 0.0 0.0 2.4 0.1 0.0 0.0Gammarus lawrencianus 3.8 0.7 15.4 7.4 - - 0.0 0.0 0.0 0.0 0.0 0.0

Table 19. Summary of other striped searobin diet studies for the coastal Middle Atlantic Bight. (Major prey taxa ranked in relative order of importance by: mean percent

frequency of occurrence (FO), mean percent contribution to total stomach content volume (TV), mean percent contribution to total stomach content weight (TW),and mean percent contribution to total number of individual items in the stomach (TN), as available from each source.)

Year(s) No. Examined

(month(s), season(s), (size [TL cm]) or Primary Prey in Order of Importance

Source Study Area and/or frequency) age range, mean size) by FO, IRI, TDW, TN, TV, and/or TW

Marshall (1946) New Haven, CT 1942 (summer) 28 (YOY) FO -- Crangon and unidentified fish.Marshall (1946) Woods Hole, MA 1943-44 (summer) 10 (3-6) FO -- Crangon, copepods, and eggs.

11 (18-30) FO -- Neomysis, Gammarus sp., Crangon, and fish

(northern searobin, unidentified flounder, and others).

Kimmel (1973) Mouth of Chesapeake 1971-72 7 (10-14) TV -- Neomysis and Crangon.

Bay (spring/summer)
Page 53

Richards et al. (1979) Long Island Sound, CT 1971-72 (spring/fall) 124 (4-14) TN -- Neomysis, copepods (Temora sp. and Labidocera sp.),

and Crangon.

1976-77 (same) 57 (12-33) TN -- Crangon and Neomysis.

Mann (1974) Long Island Sound, NY 1973 (spring-fall) 533 (YOY) TN -- Neomysis and Crangon.

390 (1-9) TN -- Crabs, Crangon, juvenile fish (scup, northern and

striped searobins, windowpane, winter flounder, bay anchovy,Menidia, and northern pipefish), and Neomysis.

Carlson (1991) New Haven Harbor, CT 1989-90 (monthly) 15 (32-39) TW -- Ovalipes ocellatus, juvenile winter flounder, and

juvenile Atlantic rock crab.FO -- O. ocellatus, Atlantic rock crab, and juvenile winter

flounder.

Manderson et al. Sandy Hook Bay, NJ 1998 (Jun-Oct) 77 (18-37, mean = 28) TW -- Crangon, mysids (Neomysis), juvenile winter flounder,

(1999) amphipods, and juvenile O. ocellatus.

FO -- Crangon, mysids, juvenile winter flounder, amphipods,and O. ocellatus.
Page 54Table 20. Summary of Hudson-Raritan Estuary northern searobin diet by sampling period during 1996-97, expressed as the percent frequency of occurrence of dominant prey

or items (FO), and as the mean percent contribution of dominant prey or items to total stomach content volume (TV). (n = sample size; ES = percent of emptystomachs.)

Sampling Period

Jul 96 Oct 96 Jan 97 Apr 97 Aug 97 Nov 97

(n = 0; ES = N/A) (n = 2; ES = 0) (n = 0; ES = N/A) (n = 10; ES = 40) (n = 87; ES = 5) (n = 4; ES = 0)

Prey FO TV FO TV FO TV FO TV FO TV FO TV

Crangon septemspinosa - - 50.0 16.7 - - 10.0 16.7 77.0 37.0 50.0 33.3

Neomysis americana - - 100.0 33.3 - - 10.0 16.7 16.1 4.0 50.0 16.7Gammarus lawrencianus - - 100.0 33.3 - - 0.0 0.0 5.7 3.6 25.0 8.3Unidentified crustaceans - - 0.0 0.0 - - 30.0 25.0 1.1 0.1 0.0 0.0Juvenile Atlantic rock crab - - 0.0 0.0 - - 0.0 0.0 37.9 23.6 0.0 0.0Unciola sp. - - 50.0 16.7 - - 0.0 0.0 2.3 1.0 0.0 0.0Juvenile Ovalipes ocellatus - - 0.0 0.0 - - 0.0 0.0 9.2 6.2 0.0 0.0Unidentified amphipods - - 0.0 0.0 - - 0.0 0.0 2.3 0.5 25.0 8.3Unidentified juvenile fish - - 0.0 0.0 - - 0.0 0.0 4.6 1.6 25.0 25.0Coal pebbles - - 0.0 0.0 - - 0.0 0.0 24.1 6.8 0.0 0.0

Table 21. Summary of other northern searobin diet studies for the coastal Middle Atlantic Bight. (Major prey taxa ranked in relative order of importance by: mean percent

frequency of occurrence (FO), mean percent contribution to total stomach content volume (TV), mean percent contribution to total stomach content weight (TW),and mean percent contribution to total number of individual items in the stomach (TN), as available from each source.)

Year(s) No. Examined

(month(s), season(s), (size [TL cm]) or Primary Prey in Order of Importance

Source Study Area and/or frequency) age range, mean size) by FO, IRI, TDW, TN, TV, and/or TW

Linton (1901) Woods Hole, MA 1899-1900 (summer) Unknown FO? -- Herring, softshells, Palaemonetes sp., amphipods,

squid, “clams,” polychaetes, and juvenile winter flounder.

Marshall (1946) Woods Hole , MA 1943 (summer) 42 (2-7) FO -- Amphipods (Gammarus locusta), Crangon,

copepods, and Neomysis.

38 (>13) FO -- Crangon, G. locusta, Neomysis, Ampelisca sp., and

polychaetes.

Marshall (1946) Block Island Sound 1943 (summer) 13 (>17) FO -- Ampelisca sp., G. locusta, Unciola irrorata, Cerapus

tubularis, and polychaetes.
Page 55

Marshall (1946) Montauk Point, NY 1943 (spring) 12 (18-25) FO -- Monoculodes edwardsi, Neomysis, cumaceans, and

Crangon.

Smith (1950) Block Island Sound 1948-49 (seasonally) Unknown TV? -- Leptocheirus pinguis, U. irrorata, and Crangon.

Richards (1963) Long Island Sound 1956-57 (seasonally) Sand station - 103 FO? -- Copepods, amphipods, Neomysis, Heteromysis

(2-16) formosa, Pagurus sp., Crangon, bay anchovy, and Cyathura

sp.

Mud station - 23 FO? -- Copepods, amphipods (Unciola sp.), Neomysis, and

(2-10) cumaceans.

Kimmel (1973) Mouth of 1971-72 (spring- 47 (2-12) TV -- Neomysis, Crangon, and A. macrocephala.

Chesapeake Bay summer)

Richards et al. (1979) Long Island Sound 1971-73 (spring-fall) 120 (4-12) TN -- Neomysis, copepods (Acartia sp., Pseudodiaptomus

coronatus, and Temora sp.), and Crangon.

1976-77 25 (10-30) TN -- Amphipods, isopods, crabs, Crangon, and Neomysis.

Mann (1974) Long Island Sound, 1973 (May-Dec) 419 (~10-29) TN -- Crangon, crabs, bivalve mollusks (Ensis directus),

NY amphipods (Unciola sp.), isopods, cumaceans, and

copepods.

Allen et al. (1978) Hereford Inlet, NJ 1973-77 (biweekly/ 107 (4-21) FO -- Amphipods, Crangon, mysids, crabs, unidentified

monthly) fish, polychaetes, crab larvae, and Pagurus sp.

Homer and Boynton Chesapeake Bay, MD 1977 (May/Sep) Unknown (~5-18) TW -- Juvenile weakfish, Nereis succinea, amphipods,

(1978) polychaetes, and anemones.
Page 56Table 22. Summary of Hudson-Raritan Estuary striped bass diet by sampling period during 1996-97, expressed as the percent frequency of occurrence of dominant prey or items (FO), and as the mean percent contribution of dominant prey or items to total stomach content volume (TV). (n = sample size; ES = percent of empty stomachs.)

Sampling Period

Jul 96 Oct 96 Jan 97 Apr 97 Aug 97 Nov 97

(n = 0; ES = N/A) (n = 3; ES = 100) (n = 24; ES = 37) (n = 32; ES = 25) (n = 0; ES = N/A) (n = 22; ES = 9)

Prey FO TV FO TV FO TV FO TV FO TV FO TV

Crangon septemspinosa - - 0.0 0.0 58.3 100.0 75.9 36.9 - - 54.5 15.0

Neomysis americana - - 0.0 0.0 0.0 0.0 31.0 16.6 - - 4.5 <0.1 Unidentified fish remains - - 0.0 0.0 0.0 0.0 3.4 0.3 - - 36.4 1.9

Smallmouth flounder - - 0.0 0.0 0.0 0.0 0.0 0.0 - - 18.2 12.3

Palaemonetes sp. - - 0.0 0.0 0.0 0.0 0.0 0.0 - - 13.6 5.1

Ampelisca abdita - - 0.0 0.0 0.0 0.0 17.2 6.6 - - 0.0 0.0 Juvenile red hake - - 0.0 0.0 0.0 0.0 0.0 0.0 - - 13.6 10.6 Unidentified goby - - 0.0 0.0 0.0 0.0 0.0 0.0 - - 9.1 0.8 Juvenile silver hake - - 0.0 0.0 0.0 0.0 0.0 0.0 - - 13.6 3.4

Juvenile Ovalipes ocellatus - - 0.0 0.0 0.0 0.0 0.0 0.0 - - 13.6 4.2

Nassarius trivittatus - - 0.0 0.0 0.0 0.0 3.4 2.9 - - 4.5 5.9 Northern searobin - - 0.0 0.0 0.0 0.0 3.4 2.4 - - 4.5 5.9

Table 23. Summary of other striped bass diet studies for the coastal Middle Atlantic Bight. (Major prey taxa ranked in relative order of importance by: mean percent frequency

of occurrence (FO), mean percent contribution to total stomach content volume (TV), mean percent contribution to total stomach content weight (TW), and meanpercent contribution to total number of individual items in the stomach (TN), as available from each source.)

Year(s) No. Examined

(month(s), season(s), (size [TL cm]) or Primary Prey in Order of Importance

Source Study Area and/or frequency) age range, mean size) by FO, IRI, TDW, TN, TV, and/or TW

Curran and Reis (1937) Lower Hudson River, 1936 (unknown) Unknown (“small”) FO? -- Gammarus fasciatus, chironomids, and copepods.

NY

Merriman (1941) Connecticut coast 1936-37 (Apr-Nov) 550 (“juveniles”) FO? -- Menidia and juvenile Atlantic menhaden.

Hollis (1952) Chesapeake Bay 1936-37 (seasonally) 1,736 (~14-74) TN -- Bay anchovy, Atlantic menhaden, spot, croaker,

herring, unidentified fish, cladocerans, isopods, Crangon,and Palaemonetes sp.

de Sylva et al. (1962) Delaware Bay 1958-60 (Mar-Dec) 279 (4-31) FO -- Neomysis, Gammarus sp., Crangon, Palaemonetes

sp., other crustaceans, Limulus polyphemus eggs and larvae,

and unidentified fish.
Page 57

Schaefer (1970) Southern Long Island 1964 (Apr-Nov) 61 (28-40) TV -- Amphipods (Gammarus sp. and haustorids),

surf zone Neomysis, Atlantic surfclam, and bay anchovy.

183 (41-60) TV -- Amphipods (Gammarus sp. and haustorids), bay

anchovy, Menidia, and scup.

123 (61-94) TV -- Amphipods (Gammarus sp. and haustorids), red hake,

Ovalipes ocellatus, scup, bay anchovy, juvenile tautog,northern puffer, and mullet.

Markle and Grant York River, VA 1967 (Jul-Oct) 297 (3-15) TV -- Fish (goby, Menidia, and shiners), Palaemonetes sp.,

(1970) Crangon, Neomysis, and polychaetes.

Festa (1979) Little Egg Harbor, NJ 1972-77 (fall) 11 (43-58) TV -- Sand lance, weakfish, unidentified fish, and Crangon.

FO -- Sand lance, unidentified fish, and Crangon.

Gardinier and Hoff Middle Hudson River 1974 (Apr-Nov) 546 (<15) FO -- Gammarus sp., calanoid copepods, and juvenile fish(1982) Estuary (Atlantic tomcod).

 (15-28) FO -- Fish (herring, Atlantic tomcod, bay anchovy, banded

killifish, and mummichog).

1976-77 (Apr-May) 380 (>20) FO -- Fish (herring, bay anchovy, shiners, and killifish),

detritus, and “invertebrates.”

Boynton et al. Potomac River, 1976 (biweekly, Jun- 703 (2.5-10) TN -- Fish, mysids, and polychaetes.

(1981) VA/MD Sep, higher salinity area)Homer and Chesapeake Bay, MD 1977 (spring) 19 (10-15) TW -- Crangon, juvenile Atlantic menhaden, Nereis

Boynton (1978) succinea, bay anchovy, and juvenile spot.

Lawler, Matusky & Hudson River, NY 1979-80 (seasonally) 239 (mean = 18.50) TV -- Gammarus sp., Crangon, Paleomonetes sp., and

Skelly Engineers (1980) Neomysis.

Fay et al. (1982) Hudson River Unknown (<7.5) FO? -- Gammarus sp., calanoid copepods, and insect larva.

 (7.6-12.5) FO? -- Gammarus sp. and calanoid copepods.

 (13-20) FO? -- Atlantic tomcod.

Fay et al. (1982) Delaware River Unknown (5-10) FO? -- Neomysis and Crangon.

Fay et al. (1982) Chesapeake Bay Unknown (7-15) FO? -- Naked goby and Palaemonetes sp.

 (10-27) FO? -- Bay anchovy.

Hartman and Chesapeake Bay 1990-92 (bimonthly) (age 0) TW -- Juvenile Atlantic menhaden, spot, and other fish.

Brandt (1995) (age 1) TW -- “Invertebrates,” Atlantic menhaden, bay anchovy,

and Gammarus sp.

 (age 2) TW -- Atlantic menhaden, spot, and other fish.
Page 58Table 24. Summary of Hudson-Raritan Estuary clearnose skate diet by sampling period during 1996-97, expressed as the percent frequency of occurrence of dominant prey or

items (FO), and as the mean percent contribution of dominant prey or items to total stomach content volume (TV). (n = sample size; ES = percent of empty stomachs.)

Sampling Period

Jul 96 Oct 96 Jan 97 Apr 97 Aug 97 Nov 97

(n = 27; ES = 0) (n = 13; ES = 0) (n = 0; ES = N/A) (n = 0; ES = N/A) (n = 30; ES = 0) (n = 1; ES = 0)

Prey FO TV FO TV FO TV FO TV FO TV FO TV

Crangon septemspinosa 78.8 25.1 61.5 29.2 ---- 36.7 15.1 100.0 33.3Atlantic rock crab 60.6 37.2 7.7 1.9 ---- 73.4 25.7 0.0 0.0Unidentified fish remains 9.1 2.3 46.2 17.7 ---- 20.0 10.1 100.0 33.3Ovalipes ocellatus 6.1 2.5 15.4 9.8 ---- 13.3 7.6 100.0 33.3Juvenile conger eel 0.0 0.0 15.4 11.3 - - - - 0.0 0.0 0.0 0.0Juvenile winter flounder 3.0 2.9 7.7 11.3 - - - - 0.0 0.0 0.0 0.0Unidentified juvenile flounder 0.0 0.0 0.0 0.0 ---- 23.3 13.3 0.0 0.0Unidentified organic matter 6.1 1.7 23.1 6.8 - - - - 3.3 <0.1 0.0 0.0Xanthid crabs 12.1 6.4 0.0 0.0 - - - - 6.7 2.2 0.0 0.0Juvenile windowpane 6.1 4.7 0.0 0.0 - - - - 3.3 1.7 0.0 0.0Ensis directus 6.1 6.5 0.0 0.0 - - - - 6.7 3.3 0.0 0.0
Page 59

Table 25. Summary of other clearnose skate diet studies for the coastal Middle Atlantic Bight. (Major prey taxa ranked in relative order of importance by: mean percent

frequency of occurrence (FO), and mean percent contribution to total stomach content volume (TV), as available from each source.)

Year(s) No. Examined

(month(s), season(s), (size [TL cm]) or Primary Prey in Order of Importance

Source Study Area and/or frequency) age range, mean size) by FO, IRI, TDW, TN, TV, and/or TW

Fitz and Daiber (1963) Delaware Bay 1954-56 (seasonally) 363 (mean = 56) FO -- Crangon, Ensis directus, xanthid crabs

(Dyspanopeus), Libinia dubia, Pagurus sp.,Neomysis, and fish (weakfish, windowpane, bayanchovy, and others).

Kimmel (1973) Mouth of Chesapeake 1971-72 (monthly) 28 (19-44) TV -- Crangon, E. directus, Solen sp.,

Bay Upogebia sp., Neomysis, and unidentified fish.

FO — Crangon, Neomysis, unidentified fish,

Upogebia sp., and Neomysis.

Steimle (unpubl. data) Delaware Bay 1989-94 (May/Jun 96 (28.0-78.0, mean TV -- Atlantic rock crab, E. directus,

and Aug/Sep) = 58.0) unidentified fish and crustaceans, Pagurus sp.,

Dyspanopeus sayi, and Crangon.
Page 60Table 26. Summary of Hudson-Raritan Estuary bluefish diet by sampling period during 1996-97, expressed as the percent frequency of occurrence of dominant prey or items

(FO), and as the mean percent contribution of dominant prey or items to total stomach content volume (TV). (n = sample size; ES = percent of empty stomachs.)

Sampling Period

Jul 96 Oct 96 Jan 97 Apr 97 Aug 97 Nov 97

(n = 0; ES = N/A) (n = 1; ES = 0) (n = 0; ES = N/A) (n = 0; ES = N/A) (n = 62; ES = 0) (n = 0; ES = N/A)

Prey FO TV FO TV FO TV FO TV FO TV FO TV

Unidentified fish remains - - 0.0 0.0 ---- 41.9 41.4 - -

Crangon septemspinosa - - 0.0 0.0 - - - - 24.2 22.2 - -Neomysis americana - - 0.0 0.0 - - - - 21.0 17.3 - -Unidentified organic matter - - 0.0 0.0 - - - - 9.7 3.7 - -Juvenile butterfish - - 0.0 0.0 - - - - 1.6 9.3 - -Anchoa sp. - - 100.0 100.0 - - - - 1.6 1.9 - -

Table 27. Summary of other bluefish diet studies for the coastal Middle Atlantic Bight. (Major prey taxa ranked in relative order of importance by: mean percent frequency

of occurrence (FO), mean percent contribution to total stomach content volume (TV), and mean percent contribution to total stomach content weight (TW), asavailable from each source.)

Year(s) No. Examined

(month(s), season(s), (size [TL cm]) or Primary Prey in Order of Importance

Source Study Area and/or frequency) age range, mean size) by FO, IRI, TDW, TN, TV, and/or TW

Peck (1896) Woods Hole, MA 1893 (Jul-Aug) 38 (?) FO -- Atlantic menhaden, butterfish, herring, squid, and

YOY fish.

Breder (1922a) Atlantic City, NJ 1920 (Aug) 31 (14-21) TV -- Butterfish, juvenile spot, unidentified fish, and

plant material.

Grant (1962) Indian River, DE 1956-58 (summer) 262 (3-24) TV -- Mummichog, juvenile Atlantic menhaden, Menidia,

bay anchovy, and unidentified fish.FO -- Unidentified fish, juvenile Atlantic menhaden,Menidia, mummichog, and polychaetes.

de Sylva et al. (1962) Delaware Bay shore 1958-60 (summer) 152 (4-23) FO -- Unidentified fish, Menidia, Neomysis, Crangon,

herring, and bay anchovy.
Page 61

Festa (1979) Little Egg Harbor, NJ 1972-77 (summer/fall) 16 (6-10) TV -- Palaemonetes sp., unidentified fish, Nereis sp.,

Nephtys sp., and Menidia.

FO -- Unidentified fish, crab larvae, and Nereis sp.

62 (11-20) TV -- Unidentified fish, mummichog, Menidia, juvenile

blue crab, and bay anchovy.FO -- Unidentified fish, Crangon, detritus, bay anchovy,

Menidia, and Neomysis.

Allen et al. (1978) Hereford Inlet, NJ 1973-77 (biweekly/ 301 (3-18) FO -- Unidentified fish, Crangon, Menidia, crab larvae,

monthly) juvenile spot, and shrimp larvae.

Bason et al. (1976) Chesapeake and 1975 (spring-summer) 201 (5-33) TV -- Bay anchovy, juvenile Atlantic menhaden, juvenile

Delaware Canal, DE weakfish, unidentified fish, Menidia, and juvenile

bluefish.

Lynch et al. (1977) Raritan River and Bay 1976-77 (Apr-Mar) 10 (3-22) FO? -- Mummichog, bay anchovy, Menidia, Crangon,

Palaemonetes sp., and fish.

39 (39+) FO? -- Atlantic menhaden, spot, bay anchovy, and

Crangon.

Friedland et al. (1988) Sandy Hook Bay, NJ 1981-84 (spring- 1,079 (9-18) TW -- Bay anchovy, Crangon, Menidia, Neomysis,

summer) striped killifish, and Palaemonetes sp.

FO -- Crangon, Neomysis, bay anchovy, Menidia,Palaemonetes sp., and striped killifish.

Scarlett and Giust (1989) Manasquan River, NJ 1984-86 (summer) 211 (9-29, mean TV -- Bay anchovy, Menidia, Crangon, unidentified fish,

= ~15) Gammarus sp., and juvenile weakfish.

Juanes et al. (1993) Hudson River, NY 1989 (Jul-Oct) 374 (YOY) TW -- Atlantic tomcod, unidentified fish, juvenile striped

bass, juvenile bluefish, and bay anchovy.

Juanes and Conover (1994) Great South Bay, NY 1989 (Jun-Jul) 256 (YOY) TW (Jun) -- Menidia, “shrimp,” other fish, and bay

anchovy.TW (Jul) -- bay anchovy, crustaceans, other fish, andMenidia.

Hartman and Brandt (1995) Chesapeake Bay 1990-92 (bimonthly) 100 (age 0) TW -- Bay anchovy, juvenile Atlantic menhaden,

Menidia.

132 (age 1) TW -- Spot, juvenile Atlantic menhaden, and other fish.
Page 62Table 28. Summary of Hudson-Raritan Estuary winter skate diet by sampling period during 1996-97, expressed as the percent frequency of occurrence of dominant prey or

items (FO), and as the mean percent contribution of dominant prey or items to total stomach content volume (TV). (n = sample size; ES = percent of empty stomachs.)

Sampling Period

Jul 96 Oct 96 Jan 97 Apr 97 Aug 97 Nov 97

(n = 0; ES = N/A) (n = 5; ES = 20) (n = 19; ES = 0) (n = 24; ES = 0) (n = 0; ES = N/A) (n = 9; ES = 0)

Prey FO TV FO TV FO TV FO TV FO TV FO TV

Crangon septemspinosa - - 40.0 14.9 100.0 44.8 95.8 42.7 - - 77.8 28.2

Unidentified fish remains - - 20.0 9.0 15.8 5.6 4.2 1.2 - - 22.2 9.8Unidentified flounder - - 20.0 7.5 5.3 1.1 8.3 5.0 - - 11.1 3.3Atlantic rock crab - - 40.0 11.9 36.8 25.0 8.3 2.7 - - 0.0 0.0Blue mussel - - 20.0 4.5 0.0 0.0 4.1 1.3 - - 11.1 3.6Juvenile winter flounder - - 0.0 0.0 10.5 4.5 4.1 1.3 - - 33.3 14.9Ovalipes ocellatus - - 40.0 16.4 0.0 0.0 8.3 3.8 - - 11.1 3.9Sand lance - - 60.0 19.4 0.0 0.0 0.0 0.0 - - 33.3 12.8
Page 63

Table 29. Summary of other winter skate diet studies for the coastal Middle Atlantic Bight. (Major prey taxa ranked in relative order of importance by: mean percent

contribution to total stomach content weight (TW), and an index of relative importance (IRI), as available from each source.)

Year(s) No. Examined

(month(s), season(s), (size [TL cm]) or Primary Prey in Order of Importance

Source Study Area and/or frequency) age range, mean size) by FO, IRI, TDW, TN, TV, and/or TW

Smith (1950) Block Island Sound 1848-49 (seasonally) Unknown (?) TW -- “Nekton” (fish and squid), Leptocheirus

pinguis, and Nephtys incisa.

Vinogradov (1984) Southern New England 1968-74 (various) 52 (unknown) TW -- Crustaceans, polychaetes, and fish.McEachlan et al. Middle Atlantic Bight 1969-70 (seasonally) 441 (mode = ~60) IRI -- Amphipods (L. pinguis), polychaetes (N. incisa,

(1976) shelf Nereis sp., Pectinaria gouldii, and maldanids), fish (sand

lance), and decapod crustaceans (Crangon).
Page 64Table 30. Summary of Hudson-Raritan Estuary black sea bass diet by sampling period during 1996-97, expressed as the percent frequency of occurrence of dominant prey or

items (FO), and as the mean percent contribution of dominant prey or items to total stomach content volume (TV). (n = sample size; ES = percent of empty stomachs.)

Sampling Period

Jul 96 Oct 96 Jan 97 Apr 97 Aug 97 Nov 97

(n = 1; ES = 0) (n = 3; ES = 0) (n = 0; ES = N/A) (n = 3; ES = 33) (n = 46; ES = 11) (n = 2; ES = 0)

Prey FO TV FO TV FO TV FO TV FO TV FO TV

Crangon septemspinosa 100.0 100.0 66.7 50.0 - - 0.0 0.0 43.5 24.0 0.0 0.0Neomysis americana 0.0 0.0 0.0 0.0 - - 33.3 50.0 26.1 13.9 50.0 12.5Juvenile Atlantic rock crab 0.0 0.0 33.3 2.9 - - 0.0 0.0 17.4 16.8 0.0 0.0Unidentified crustaceans 0.0 0.0 0.0 0.0 - - 33.3 50.0 4.3 3.2 0.0 0.0Corophium sp. 0.0 0.0 33.3 47.1 - - 0.0 0.0 2.2 0.1 50.0 12.5Xanthid crabs 0.0 0.0 0.0 0.0 - - 0.0 0.0 8.7 10.3 0.0 0.0Ampelisca abdita 0.0 0.0 0.0 0.0 - - 0.0 0.0 15.2 1.0 0.0 0.0Juvenile Atlantic menhaden 0.0 0.0 0.0 0.0 - - 0.0 0.0 0.0 0.0 50.0 75.0Unidentified organic matter 0.0 0.0 0.0 0.0 - - 0.0 0.0 21.7 3.5 0.0 0.0

Table 31. Summary of other juvenile black sea bass diet studies for the coastal Middle Atlantic Bight. (Major prey taxa ranked in relative order of importance by: mean percent

frequency of occurrence (FO), and mean percent contribution to total stomach content volume (TV), as available from each source.) [Note: there are other studies ofthe diet of black sea bass, but they were not included in this table because they were concerned with larger fish (e.g., Steimle and Figley 1996), or because the size or

life stage discussed was ambiguous, or because only a small sample (e.g., less than 10 fish) was examined (e.g., Steimle and Ogren 1982).]

Year(s) No. Examined

(month(s), season(s), (size [TL cm]) or Primary Prey in Order of Importance

Source Study Area and/or frequency) age range, mean size) by FO, IRI, TDW, TN, TV, and/or TW

Peck (1896) Woods Hole, MA 1893 (Jul-Aug) 40 (unknown) FO -- Carcinus maenas, Ovalipes ocellatus, amphipods,

small fish (mostly sculpins), gastropods, American lobster,and Pagurus sp.

Richards (1963) Long Island Sound - 1956-58 (Sep-Oct) 28 (2-4) FO? -- Caprellid amphipods, other amphipods (Stenothoe

sand station sp., Corophium sp., and Ericthonius sp.), hydroids,

copepods, mysids, and Pagurus sp.

Kimmel (1973) Mouth of 1971-72 (spring/ 48 (3-15) TV -- Neomysis, unidentified polychaetes, xanthid (mud)

Chesapeake Bay summer) crabs, and amphipods.
Page 65

Allen et al. (1978) Hereford Inlet, NJ 1973-77 (biweekly/ 201 (4-20) FO -- Mysids, Crangon, detritus, amphipods, crabs,

monthly) polychaetes, and unidentified fish.

Chee (1977) Coastal Virginia 1975-76 (monthly) 26 (<15) TV -- Pagurus sp., O. ocellatus, and Atlantic rock crab.

Festa (1979) Little Egg Harbor, NJ 1976 (summer) 17 (6-21) TV -- Mud crabs (Neopanope sp.), O. ocellatus, blue crab,

Palaemonetes sp., and Crangon.FO -- Mud crabs, Crangon, isopod (Idotea sp.), amphipods

(Elasmopus sp. and Ampelisca sp.), Palaemonetes sp., blue

crab, and Neomysis.

Steimle (unpubl. Delaware Bay 1989-94 (May/Jun 185 (11.0-39.0, mean TV -- Dyspanopeus sayi, Atlantic rock crab, Neomysis, blue

data) and Aug/Sep) = 24.0) mussel, Pagurus sp., Crangon, and Ensis directus.

FO -- Neomysis, Atlantic rock crab, Dyspanopeus sayi, blue

mussel, and Pagurus sp.
Page 66Table 32. Summary of Hudson-Raritan Estuary spot diet by sampling period during 1996-97, expressed as the percent frequency of occurrence of dominant prey or items (FO),

and as the mean percent contribution of dominant prey or items to total stomach content volume (TV). (n = sample size; ES = percent of empty stomachs.)

Sampling Period

Jul 96 Oct 96 Jan 97 Apr 97 Aug 97 Nov 97

(n = 0; ES = N/A) (n = 40; ES = 43) (n = 6; ES = 83) (n = 0; ES = N/A) (n = 1; ES = 0) (n = 0; ES = N/A)

Prey FO TV FO TV FO TV FO TV FO TV FO TV

Ampelisca abdita - - 30.0 24.2 0.0 0.0 - - 100.0 50.0 - -Unidentified organic matter - - 32.5 8.2 0.0 0.0 - - 0.0 0.0 - -Neomysis americana - - 17.5 11.5 16.7 100.0 - - 0.0 0.0 - -Crangon septemspinosa - - 20.0 11.5 0.0 0.0 - - 0.0 0.0 - -Unidentified copepods - - 0.0 0.0 0.0 0.0 - - 100.0 50.0 - -Pseudodiaptomus sp. - - 12.5 18.6 0.0 0.0 - - 0.0 0.0 - -

Table 33. Summary of other spot diet studies for the coastal Middle Atlantic Bight. (Major prey taxa ranked in relative order of importance by: mean percent frequency of

occurrence (FO), mean percent contribution to total stomach content volume (TV), and mean percent contribution to total stomach content weight (TW), asavailable from each source.)

Year(s) No. Examined

(month(s), season(s), (size [TL cm]) or Primary Prey in Order of Importance

Source Study Area and/or frequency) age range, mean size) by FO, IRI, TDW, TN, TV, and/or TW

Van Engle and York River, VA Unknown 162 (juveniles) TV -- Polychaetes, amphipods, clam siphons, and

Joseph (1968) cumaceans.

FO -- Harpacticoid copepods, Gammarus sp.,nematodes, cumaceans, polychaetes, and Neomysis.

Thomas (1971) Delaware River 1970 (Oct) 17 (unknown) FO -- Copepods, Gammarus sp., Monoculodes sp.,

ostracods, detritus, and Neomysis.

Kimmel (1973) Mouth of Chesapeake 1971-72 13 (YOY, <10) TV -- Detritus and harpacticoid and calanoid copepods.

Bay (spring/summer) 35 (10-16) TV -- Detritus, Ampelisca macrocephalus, harpacticoid

copepods, and Nereis.

Festa (1978) Little Egg Harbor, NJ 1972-77 (summer/fall) 48 (3-10) TV -- Copepods and Ampelisca sp.

FO -- Copepods and amphipods.

44 (11-19) TV -- Polychaetes, amphipods, copepods, and Neomysis.

FO -- Copepods, detritus, polychaetes, and Ampelisca sp.
Page 67

Chao and Musick York River, VA 1973 (summer) 77 (7-20) FO -- Pectinaria gouldii, Glycinde solitaria, nematodes,

(1977) Neomysis, Nereis succinea, clams, detritus, copepods, and

cumaceans.

Allen et al. (1978) Hereford Inlet, NJ 1973-77 (biweekly/ 722 (2-17) FO -- Copepods, polychaetes, amphipods, detritus,

monthly) isopods, and mysids.

Homer and Boynton Chesapeake Bay, MD 1977 (May-Dec) 1,400 (~2-15) TW -- N. succinea, Scolecolepides viridis, Paraprionospio(1978) pinnata, Eteone sp., juvenile Macoma sp., and softshells.

Scarlett and Giust (1989) Manasquan River, NJ 1984-85 (summer) 8 (17-18) TV -- Detritus, Crangon, Neomysis, and Ampelisca sp.

Pihl et al. (1992) York River, VA 1989 (Jun-Sep ) 649 (5-20) TW -- Polychaetes (esp. P. pinnata), amphipods, and

mysids.

Timmons (1995) Indian River, DE 1994 (July) 20 (mean = 9) TV -- Detritus, polychaetes, and mysids.
Page 68Table 34. Summary of Hudson-Raritan Estuary American lobster diet by sampling period during 1996-97, expressed as the percent frequency of occurrence of dominant prey

or items (FO), and as the mean percent contribution of dominant prey or items to total stomach content volume (TV). (n = sample size; ES = percent of emptystomachs.)

Sampling Period

Jul 96 Oct 96 Jan 97 Apr 97 Aug 97 Nov 97

(n = 0; ES = N/A) (n = 6; ES = 0) (n = 6; ES = 0) (n = 10; ES = 0) (n = 13; ES = 0) (n = 12; ES = 0)

Prey FO TV FO TV FO TV FO TV FO TV FO TV

Atlantic rock crab - - 0.0 0.0 67.7 16.7 50.0 14.9 42.1 18.4 41.6 18.3Nucula sp. - - 0.0 0.0 50.0 15.2 30.0 13.0 30.8 7.0 16.7 0.4Pagurus sp. - - 16.7 4.7 16.7 8.7 10.0 1.3 38.5 13.2 33.3 3.3

Human artifacts - - 16.7 4.8 16.7 3.6 20.0 1.9 0.0 0.0 16.7 0.4Shell hash - - 50.0 23.8 16.7 3.6 10.0 0.6 15.4 4.4 0.0 0.0Ovalipes ocellatus - - 50.0 23.8 0.0 0.0 0.0 0.0 0.0 0.0 8.3 1.7Crangon septemspinosa - - 0.0 0.0 33.3 10.1 20.0 12.3 38.5 15.5 8.3 0.3Unidentified hydroids - - 16.7 4.8 33.3 9.4 10.0 1.3 7.7 0.3 8.3 5.7Sand - - 16.7 4.8 0.0 0.0 0.0 0.0 38.5 7.9 83.3 15.6Unidentified organic matter - - 16.7 4.8 66.7 15.9 40.0 11.7 15.4 6.4 0.0 0.0Unidentified crab fragments - - 0.0 0.0 16.7 3.6 40.0 11.7 7.7 0.9 8.3 0.1Unidentified fish remains - - 0.0 0.0 16.7 5.8 20.0 7.8 23.1 6.7 0.0 0.0Blue mussel - - 0.0 0.0 0.0 0.0 10.0 3.9 7.7 3.5 16.7 6.4Skate egg case fragments - - 0.0 0.0 0.0 0.0 20.0 9.7 0.0 0.0 8.3 4.7Hair and fibers - - 0.0 0.0 16.7 1.4 0.0 0.0 46.2 5.0 0.0 0.0Xanthid crabs - - 0.0 0.0 0.0 0.0 0.0 0.0 7.7 3.5 66.7 10.2
Page 69

Table 35. Summary of other American lobster diet studies for the coastal Middle Atlantic Bight. (Major prey taxa ranked in relative order of importance by mean percent

frequency of occurrence (FO). Size is carapace length (CL), based on eye socket.)

Year(s) No. Examined

(month(s), season(s), (size [TL cm]) or Primary Prey in Order of Importance

Source Study Area and/or frequency) age range, mean size) by FO, IRI, TDW, TN, TV, and/or TW

Weiss (1970) Eastern Long Island 1967-69 (monthly) 557 (1.7-9.7, mean = FO -- Atlantic rock crab, unidentified fish, Lacuna vincta,

Sound 6.3) Nereis sp., blue mussel, Astyris (Mitrella) lunata, and

Zostera marina.

Steimle and New York Bight apex 1982-85 (seasonally) 33 (not noted) FO -- Atlantic rock crab, Nucula, Ovalipes ocelatus,

Terranova (1991) Pherusa affinis, juvenile American lobster, fish, squid,

and unidentified polychaetes.

Steimle (1994) New York Bight apex 1986-89 (bimonthly) 935 (5-13.5) FO -- Unidentified fish, Atlantic rock crab, P. affinis, N.

proxima, and Asterias sp.

Steimle and Figley Central New Jersey 1996-97 (Jul-Sep) 99 (6.3-15.0, mean = FO -- Crab fragments (mostly Atlantic rock crab), Euspira

(unpubl. data) coast artificial reef 8.9) sp. operculums, shell hash, skate egg case fragments,

Pagurus sp., unidentified organic matter, Mytilus

fragments, hydroids, sand, artifacts (e.g., tire rubberparticles), and Asterias sp.
Page 70Table 36. Summary of Hudson-Raritan Estuary tautog diet by sampling period during 1996-97, expressed as the percent frequency of occurrence of dominant prey or items

(FO), and as the mean percent contribution of dominant prey or items to total stomach content volume (TV). (n = sample size; ES = percent of empty stomachs.)

Sampling Period

Jul 96 Oct 96 Jan 97 Apr 97 Aug 97 Nov 97

(n = 16; ES = 0) (n = 4; ES = 50) (n = 1; ES = 0) (n = 11; ES = 0) (n = 13; ES = 0) (n = 6; ES = 67)

Prey FO TV FO TV FO TV FO TV FO TV FO TV

Atlantic rock crab 8.3 8.9 25.0 40.0 100.0 100.0 45.5 29.4 46.2 31.3 16.7 93.9

Xanthid crabs 25.0 12.5 25.0 20.0 0.0 0.0 36.4 19.1 15.4 5.1 0.0 0.0Blue mussel 33.3 6.6 0.0 0.0 0.0 0.0 27.3 13.0 23.1 17.9 0.0 0.0Pagurus sp. 0.0 0.0 25.0 40.0 0.0 0.0 0.0 0.0 7.7 2.8 0.0 0.0Ensis directus 35.0 27.1 0.0 0.0 0.0 0.0 0.0 0.0 7.7 2.3 0.0 0.0Ovalipes ocellatus 16.7 8.8 0.0 0.0 0.0 0.0 9.1 4.4 0.0 0.0 0.0 0.0Crangon septemspinosa 8.3 8.9 0.0 0.0 0.0 0.0 9.1 2.2 0.0 0.0 0.0 0.0Unidentified organic matter 0.0 0.0 0.0 0.0 0.0 0.0 9.1 <0.1 30.8 10.9 0.0 0.0Juvenile blue crab 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 15.4 11.5 0.0 0.0

Table 37. Summary of other tautog diet studies for the coastal Middle Atlantic Bight. (Major prey taxa ranked in relative order of importance by: mean percent frequency of

occurrence (FO), mean percent contribution to total stomach content volume (TV),mean percent contribution to total stomach content dry weight (TDW), andmean percent contribution to total number of individual items in the stomach (TN), as available from each source.)

Year(s) No. Examined

(month(s), season(s), (size [TL cm]) or Primary Prey in Order of Importance

Source Study Area and/or frequency) age range, mean size) by FO, IRI, TDW, TN, TV, and/or TW

Steimle and Ogren New York Bight 1966-67 (Jul-Aug) 57 (23-29) TV -- Atlantic rock crab, sand dollar (Echinarachnius

(1982) parma), blue mussel, and hydroids.

Olla et al. (1974) Great South Bay, NY 1971-72 (unreported) 25 (20-46, mean = 33) TV -- “Mussels” and crustaceans (decapods and cirripeds).

Festa (1979) Little Egg Harbor, NJ 1972-77 (summer/fall) 18 (6-16) TV -- Isopods (Idotea sp.) and mud (xanthid) crabs.

FO -- Amphipods, isopods, and nematodes.

7 (20-32) TV -- Mud crabs and Ovalipes ocellatus.

FO -- Mud crabs.

Allen et al. (1978) Hereford Inlet, NJ 1973-77 (biweekly/ 28 (4-44) FO -- bivalve mollusks, amphipods, detritus, and

monthly) polychaetes.
Page 71

Grover (1982) New York - southern 1978 (Aug-Nov) 36 (3-7) TN -- Copepods and amphipods.

Long Island

Sogard (1992) Great Egg Harbor, NJ 1988-89 (summer) 63 (3-8) FO -- Copepods, amphipods, other crustaceans, and

ostracods.

Dorf (1994) Narragansett Bay, RI 1990-92 (various ) 82 (2.5-7) TDW -- Amphipods, copepods, and shrimp (Crangon and

Palaemonetes sp.).FO -- Copepods, amphipods, and flatworm.

 (7-10) TDW -- Shrimp and crabs.

FO -- Shrimp, flatworms, crabs, and detritus.

Lankford et al. (1995) Delaware Bay Unknown Unknown (age 2+) FO -- blue mussel.Steimle et al. (in review) Delaware Bay 1989-94 (May/Jun 371 (11.0-58.0, mean TV -- blue mussel, unidentified mollusks, Metridium

and Aug/Sep) = 32.0) senile, Atlantic rock crab, Ensis directus, Pagurus sp.

FO -- blue mussel, unidentified crustaceans, unidentifiedmollusks, and M. senile.

Steimle and Figley Central New Jersey 1997 (May-Aug) 68 (30.6-68.0; mean = FO -- Hydroids, blue mussel spat, Atlantic rock crab, crab

(unpubl. data) artificial reef 37.9) remains, caprellid amphipods, P. longicarpus, Euspira sp., P.

pollicaris, and gammarid amphipods.
Page 72Table 38. Summary of Hudson-Raritan Estuary smooth dogfish diet by sampling period during 1996-97, expressed as the percent frequency of occurrence of dominant prey

or items (FO), and as the mean percent contribution of dominant prey or items to total stomach content volume (TV). (n = sample size; ES = percent of emptystomachs.)

Sampling Period

Jul 96 Oct 96 Jan 97 Apr 97 Aug 97 Nov 97

(n = 16; ES = 0) (n = 9; ES = 0) (n = 0; ES = N/A) (n = 0; ES = N/A) (n = 17; ES = 0) (n = 0; ES = N/A)

Prey FO TV FO TV FO TV FO TV FO TV FO TV

Atlantic rock crab 81.3 33.2 44.4 16.6 - - - - 5.9 1.5 - -Ovalipes ocellatus 81.3 40.2 66.6 29.1 - - - - 0.0 0.0 - -Crangon septemspinosa 18.8 4.5 11.1 3.8 ---- 47.1 13.6 - -Ensis directus 25.0 8.8 11.1 0.9 ---- 23.5 6.5 - -Unidentified clam fragments 6.2 8.8 44.4 13.3 ---- 41.2 14.5 - -Xanthid crabs 31.2 11.7 0.0 0.0 ---- 17.6 6.2 - -Unidentified organic matter 0.0 0.0 55.6 25.5 - - - - 0.0 0.0 - -Unidentified fish remains 0.0 0.0 11.1 3.8 - - - - 5.9 1.9 - -Unidentified crab remains 0.0 0.0 11.1 3.3 - - - - 5.9 1.5 - -

Table 39. Summary of other smooth dogfish diet studies for the coastal Middle Atlantic Bight. (Major prey taxa ranked in relative order of importance by: mean percent

frequency of occurrence (FO), mean percent contribution to total stomach content volume (TV), and mean percent contribution to total stomach content weight(TW), as available from each source.)

Year(s) No. Examined

(month(s), season(s), (size [TL cm]) or Primary Prey in Order of Importance

Source Study Area and/or frequency) age range, mean size) by FO, IRI, TDW, TN, TV, and/or TW

Linton (1901) Woods Hole, MA ~1895 16 (unknown) FO -- Crabs (mud, Atlantic rock, and Libinia sp.), fish, squid,

and polychaetes.

Field (1907) Woods Hole, MA 1902-04 (summer) 388 (mean = ~100) FO -- Atlantic rock crab, Libinia sp., American lobster,

Atlantic menhaden, Ovalipes ocellatus, squid, Nereis sp.,

Zostera marina, and Pagurus sp.

Breder (1921) Atlantic City, NJ 1920 (Aug) 102 (30-64) FO -- Crabs, detritus, Z. marina, and fish.

Kimmel (1973) Mouth of Chesapeake 1971-72 (Dec-Sep) 6 (26-53) TV -- Upogebia sp., Ensis directus, and O. ocellatus.

Bay
Page 73

Festa (1979) Little Egg Harbor, NJ 1972-77 (summer) 12 (39-56) TV -- blue crab and juvenile weakfish.

FO -- blue crab, Palaemonetes sp., and Crangon.

Rountree and Able Little Egg Harbor, NJ 1988-90 (Apr-Nov) 85 (32-59) TW -- blue crab, unidentified crabs, Palaemonetes sp.,

(1996) Libinia sp., Crangon, O. ocellatus, and polychaetes.

Steimle (unpubl. data) Delaware Bay 1989-94 (May/Jun 110 (39-115, mean = 71) TV -- Atlantic rock crab, Pagurus sp., E. directus, and

and Aug/Sep) unidentified decapods.

FO -- Atlantic rock crab, Pagurus sp., E. directus, andunidentified decapods.

Gelsleichter et al. (1999) Mouth of Chesapeake 1980-92 64 (46-126+, TW -- Atlantic rock crab, blue crab, O. ocellatus, E. directus,

Bay mode ~ 106-115) and various fish.

FO -- Atlantic rock crab, blue crab, O. ocellatus, E. directus,and various fish.
Page 74Table 40. Summary of Hudson-Raritan Estuary silver hake diet by sampling period during 1996-97, expressed as the percent frequency of occurrence of dominant prey or items

(FO), and as the mean percent contribution of dominant prey or items to total stomach content volume (TV). (n = sample size; ES = percent of empty stomachs.)

Sampling Period

Jul 96 Oct 96 Jan 97 Apr 97 Aug 97 Nov 97

(n = 0; ES = N/A) (n = 1; ES = 0) (n = 6; ES = 0) (n = 6; ES = 0) (n = 0; ES = N/A) (n = 29; ES = 7)

Prey FO TV FO TV FO TV FO TV FO TV FO TV

Crangon septemspinosa - - 100.0 100.0 100.0 94.9 66.7 32.6 - - 24.1 11.8

Neomysis americana - - 0.0 0.0 16.7 5.1 50.0 15.2 - - 37.9 16.1Unidentified juvenile fish - - 0.0 0.0 0.0 0.0 50.0 28.3 - - 31.0 14.9Ampelisca abdita - - 0.0 0.0 0.0 0.0 16.7 10.9 - - 31.0 11.2Gammarus lawrencianus - - 0.0 0.0 0.0 0.0 16.7 10.9 - - 31.0 12.4Juvenile silver hake - - 0.0 0.0 0.0 0.0 0.0 0.0 - - 3.4 5.6Calanoid copepods - - 0.0 0.0 0.0 0.0 0.0 0.0 - - 13.7 4.3Unidentified organic matter - - 0.0 0.0 0.0 0.0 16.7 2.2 - - 6.9 1.2

Table 41. Summary of other juvenile-or-older silver hake diet studies for the coastal Middle Atlantic Bight. (Major prey taxa ranked in relative order of importance by: mean

percent frequency of occurrence (FO), mean percent contribution to total stomach content weight (TW), mean percent contribution to total stomach content dryweight (TDW), and an index of relative importance (IRI), as available from each source.)

Year(s) No. Examined

(month(s), season(s), (size [TL cm]) or Primary Prey in Order of Importance

Source Study Area and/or frequency) age range, mean size) by FO, IRI, TDW, TN, TV, and/or TW

Smith (1950) Block Island Sound 1948-49 (seasonally) Unknown TW -- Nekton (fish and squid), Crangon, and Leptocheirus

pinguis.

Richards (1963) Long Island Sound 1956-57 (seasonally) Sand station - 111 (7- FO -- Neomysis, Crangon, L. pinguis, and fish (bay

21) anchovy, sand lance, and juvenile silver hake).Mud station - 165 (6- FO -- Copepods, Neomysis, L. pinguis, Crangon, squid,

22) and bay anchovy.

Schaefer (1960) Long Branch, NJ, pier 1957-58 (Dec-Jan) 137 (~23-43) FO -- Amphipods, fish, Crangon, blueback herring,

Menidia, and mysids.TV -- Blueback herring, Menidia, juvenile silver hake, fish,and Crangon.
Page 75

Schaefer (1960) New York Bight apex 1958 (Feb-May) 201 (~23-43) FO -- Crangon, juvenile silver hake, fish, and mysids.

TV -- Juvenile silver hake, Crangon, Loligo, juvenile red

hake, blueback herring, and fish.

Steimle (1985) New York Bight apex 1968-70 (summer) 235 (7-40, mean = 27) FO -- Neomysis, Crangon, and various fish.

Vinogradov (1984) Southern New England 1968-74 (unreported) 3,233 (unreported) TW -- Juvenile silver hake, Atlantic mackerel, red hake, and

squid.

Bowman et al. (1987) Southern New England 1973-76 (spring/fall) 580 (<10) TW -- Crustaceans (Crangon, Dichelopandalus

leptocerus, Neomysis, and amphipods) and fish (sand lanceand juvenile silver hake).

 (11-20) TW -- Crangon, D. leptocerus, euphausids, Neomysis,

juvenile silver hake, and sand lance.

Sedberry (1983) New York Bight shelf 1976-77 (seasonally) 334 (1-15) IRI -- Parathemisto gaudichaudi, decapod crustaceans,

and fish (juvenile red hake and sand lance).

210 (15-25) IRI -- Decapod crustaceans, fish, and P. gaudichaudi.

Steimle and New York Bight apex 1982-85 (seasonally) 215 (not noted) FO -- Crangon, mysids, fish, D. leptocerus, juvenile silver

Terranova (1991) hake, and sand lance.Steimle (unpubl. data) New York Bight apex 1986-87 (monthly) 766 (9.0-26.2) FO -- Crangon, D. leptocerus, mysids, fish, juvenile silver

hake, and sand lance.TV -- Crangon, fish, D. leptocerus, decapod crustaceans,

juvenile silver hake, and sand lance.
Page 76Table 42. Most commonly eaten invertebrate prey in the Hudson-Raritan Estuary by the 20 most common predators examined in this survey, in approximate order of their

overall frequency of occurrence in the diets

Prey (common name) Primary Predators (also eaten by other predators)

Crangon septemspinosa Commonly eaten by all but winter flounder and tautog for which they are minor prey.(sevenspine bay shrimp or sand shrimp)Neomysis americana Eaten by all but clearnose & winter skates, tautog, smooth dogfish, and American lobster.(opossum shrimp)Gammarus lawrencianus Eaten by windowpane, scup, red, spotted & silver hakes, weakfish, northern searobin, and northern kingfish.(scud amphipod)Cancer irroratus Mostly juveniles eaten by little, clearnose & winter skates, striped & northern searobin, black sea bass, tautog,(Atlantic rock crab) smooth dogfish, grubby, American lobster, summer flounder, and red hake.Ovalipes ocellatus Juveniles mostly eaten by little, winter & clearnose skates, summer flounder, smooth dogfish, spotted hake,(lady crab or calico crab) striped searobin, and striped bass.Pagurus spp., P. longicarpus Commonly eaten by little skate, American lobster, tautog, smooth dogfish, and smallmouth flounder.

(right-handed hermit crabs, longwrist hermit)Ampelisca abdita Winter flounder, windowpane, scup, weakfish, striped searobin, black sea bass, spot, and silver hake(four-eye amphipod) commonly eat these.Pseudodiaptomus coronatus Commonly eaten by windowpane, spotted hake, and spot.(red copepod)Mytilus edulis Adults eaten by winter skate and tautog; spat eaten by other predators.(blue mussel)Corophium tuberculatum Weakfish and grubby eat them.(tube amphipod)Unciola sp. (tube amphipod) Eaten by winter flounder and northern searobin.Dyspanopeus sayi Eaten mostly by tautog, black sea bass, and smooth dogfish.(Say mud crab or xanthid crab)Ensis directus Siphons and whole clams eaten by little & clearnose skate, smooth dogfish, and tautog.(Atlantic jacknife or razor clam)
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Callinectes sapidus Minor use of juveniles by several predators.(blue crab)Palaemonetes sp., mostly P. vulgaris Found in striped bass and American lobster stomachs.

(grass shrimps, marsh grass shrimp)Dichelopandalus leptocerus Eaten by red hake, with minor use by several other predators.

(bristled longbeak shrimp or red-beaked shrimp)Asabellides oculata Minor use only by several predators.(spaghetti worm)Pherusa affinis Minor use only by several predators.(green bristlehead tube worm)Mercenaria mercenaria Only siphons eaten by winter flounder, and minor use by scup and little skate.(northern quahog or hard clam)Spisula solidissima Only siphons eaten by winter flounder, and meats by smooth dogfish.(Atlantic surfclam)Glycera sp. Minor use by winter flounder and smooth dogfish.(blood worm)
Page 78 Table 43. Habitat associations of other, less commonly eaten invertebrate prey in the Hudson-Raritan Estuary, based primarily on Caracciolo and Steimle (1983). (Scientific

and common names follow Turgeon et al. (1998) and Williams et al. (1989).)

Taxa (scientific name) Habitat Common Name and Comments

Algae (green, red, brown) Attached and loose Ulva-type fragments, branches usually found

Hydrozoans (unidentified spp.) Hard surfaces “Sea hair,” incidentally eatenNemerteans (unidentified sp.) Soft sediments, among mussels Ribbon or “tapeworm”Mollusca

Gastropoda

Crepidula fornicata Hard surfaces, marine Common Atlantic slippersnail, only juveniles usually eaten

Bivalvia

Nucula sp. Silty sediments Nut clam, tinyMytilus edulis Hard surfaces and open bottom Blue mussel, spat often within hydroidsMulinia lateralis Fine-medium estuarine sands Dwarf surfclam

Tellina agilis Silty to medium marine sands Northern dwarf-tellin

Ensis directus Fine-coarse sands Atlantic jacknife or razor clam, siphons or eaten wholeMya arenaria Soft mud-fine sands Softshell, usually only siphons nipped

Cephalopoda

Unidentified squid (Loligo?) Nektonic Fragments only found

Annelida

Polychaeta

Glycera sp. Soft to medium sands “Blood worms”

Nephtys sp. Silty to fine sands “Painted worms”Nereis sp. Silty to medium sands “Clam worms,” swarms to surface at timesAsabellides oculata Silty sands Rapid colonizer, “spaghetti worm”

Pherusa affinis Marine silty sands to mud Tube dweller

Arthropoda

Crustacea

Copepoda

Pseudodiaptomus coronatus Near bottom Bright orange color

Isopoda

Cyathura sp. Less saline, silty areas Tube dwellerEdotea sp. Silty sediments Common in detritus

 Amphipoda

Corophium tuberculatum Silty sands Tube dweller
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Ericthonius sp. Silty sands Tube dwellerUnciola sp. Various sands Tube dweller

 Decapoda

Dichelopandalus leptocerus Nektonic Bristled longbeak or bristlebeak shrimpPalaemonetes sp. In and near salt marshes Grass shrimp (mostly P. vulgaris, the marsh grass shrimp)Callinectes sapidus Estuarine, mud to sand, vegetation Blue crab, eaten at smaller sizesXanthid crabs Mud and shellfish beds Mud crabs (mostly Dyspanopeus sayi, the Say mud crab)

Stomatopoda

Squilla empusa Burrows in sediments Mantis shrimp
Page 80Table 44. Most commonly eaten fish prey in the Hudson-Raritan Estuary, in approximate order of their overall frequency of occurrence in diets

Prey Habitat Primary Predators

Prey (if definable) (minor predation by others possible)

Unidentified fish (including scales, Not definable Windowpane, summer flounder, red, spotted & silver hakes,bones and otoliths) weakfish, northern searobin, striped bass, clearnose & winter

skates, and smooth dogfish

Unidentified juvenile or small flounder Not definable Little & winter skates, summer flounder, red & spotted hakesUnidentified juvenile searobins Benthic genus, common in warmer months Minor use of juveniles by little & clearnose skates, summer

(Prionotus sp.) flounder, red & spotted hakes, striped searobin, and bluefish

Anchovy, Anchoa sp. (mostly bay Small, pelagic, schooling species, most common in Windowpane, weakfish, northern kingfish; minor use by others

anchovy) warmer months

Smallmouth flounder Small benthic species Striped bass; minor use by othersRed hake Benthic species, mostly found in the cooler months Minor use of juveniles by several speciesGoby (naked?), Gobiosoma sp. Minute benthic species, usually associated with shelter, Striped bass; minor use by skates

e.g., sponges

Juvenile silver hake Semipelagic species, commonly found in cooler months Weakfish, striped bass, larger silver hake; minor use by others

Juvenile black sea bass Structure-oriented species, often collected in redbeard Minor use by several species

sponge patches, probably also found within rocky rip-rap

Juvenile Atlantic menhaden Pelagic, schooling species, summer visitor Minor use by several speciesJuvenile windowpane Resident species, found widespread throughout the Adult red hake; minor use by other species

Estuary

Juvenile cunner Resident species usually found near structure Summer flounder; minor use by skates and red hakeSilversides, Menidia sp. Common, schooling species in shallow water during Juvenile bluefish; minor use by others

warmer months

Rock gunnel Small, resident species, associated with structure Minor use by little & clearnose skates, summer flounder, and

striped bass

Northern pipefish Small, semiresident species, often found in algal patches Minor use by several species
Page 81

Juvenile winter flounder Resident species, most common in deeper, silty areas Clearnose & winter skates; minor use by little skate

Juvenile striped searobin Benthic, warm season visitor, usually trawled in deeper Adult striped searobin; minor use by a few other species

areas

Juvenile grubby Small resident species Minor use by several speciesSand lance, Ammodytes sp. Sand-dwelling species, found in marine areas Winter skate; minor use by windowpane and little skateJuvenile weakfish Semipelagic, summer resident Minor use by summer flounder, red hake, and adult weakfishJuvenile river herring, Alosa sp. Schooling species, usually collected in cooler months Minor use by windowpane and summer flounder

Northern searobin Small, summer resident Minor use of adults and juveniles by little skate and striped

bassButterfish Small, semipelagic schooling species common in warmer Weakfish and bluefish, only

months

Juvenile American eel Prefers silty sediments Clearnose skate and spotted hakeUnidentified juvenile hake, Not definable Striped bass; minor use by clearnose skate

Urophycis sp.Other juvenile or small fish Various Winter skate, windowpane, smooth dogfish, summer flounder,infrequently preyed upon: Atlantic striped bass, and striped searobinherring, Atlantic croaker, linedseahorse, scup, conger eel, andnorthern stargazer (Astroscopusguttatus)

Page 83

Figure 1. Upper Panel: Habitat features of the Hudson-Raritan Estuary. Lower Panel: Locations of sampling strata (#s 1-9)

and blocks in the Hudson-Raritan Estuary.
Figure 2. Winter flounder distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 (Wilk et al. 1998).

Page 84
Figure 3. Windowpane distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 (Wilk et al. 1998).

Page 85
Figure 4. Little skate distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 (Wilk et al. 1998).

Page 86
Figure 5. Scup distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 (Wilk et al. 1998).

Page 87
Figure 6. Summer flounder distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 (Wilk et al. 1998).

Page 88
Figure 7. Red hake distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 (Wilk et al. 1998).

Page 89
Figure 8. Weakfish distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 (Wilk et al. 1998).

Page 90
Figure 9. Spotted hake distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 (Wilk et al. 1998).

Page 91
Figure 10. Striped searobin distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 (Wilk et al. 1998).

Page 92
Figure 11. Northern searobin distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 (Wilk et al. 1998).

Page 93
Figure 12. Striped bass distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 (Wilk et al. 1998).

Page 94
Figure 13. Clearnose skate distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 (Wilk et al. 1998).

Page 95
Figure 14. Bluefish distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 (Wilk et al. 1998).

Page 96
Figure 15. Winter skate distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 (Wilk et al. 1998).

Page 97
Figure 16. Black sea bass distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 (Wilk et al. 1998).

Page 98
Figure 17. Spot distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 (Wilk et al. 1998).

Page 99
Figure 18. American lobster distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 (Wilk et al. 1998).

Page 100
Figure 19. Tautog distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 (Wilk et al. 1998).

Page 101
Figure 20. Smooth dogfish distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 (Wilk et al. 1998).

Page 102
Figure 21. Silver hake distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 (Wilk et al. 1998).

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Figure 22. Atlantic rock crab distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 (Wilk et al. 1998).

Page 104
Figure 23. Lady crab distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 (Wilk et al. 1998).

Page 105
Figure 24. Bay anchovy distribution and relative abundance in Hudson-Raritan Estuary, 1992-97 (Wilk et al. 1998).

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