2011 NORTHEASTERN NATURALIST 18(3):291–314
Diet Composition and Feeding Habits of Common Fishes
in Long Island Bays, New York
Skyler R. Sagarese1,*, Robert M. Cerrato1, and Michael G. Frisk1
Abstract - Developing models in support of ecosystem-based management requires
knowledge of trophic dynamics of ecologically important species. A paucity of data on
these dynamics for Long Island finfish is hindering development of ecosystem models
required by recent legislation. In this study, we analyzed stomach contents of common
fishes collected from Port Jefferson Harbor, Great South Bay, and Shinnecock Bay between
May and October of 2007 and 2008. General diet composition was described by
percent by number (%N), percent by weight (%W), percent frequency of occurrence
(%O), and percent index of relative importance (%IRI) for seven species: Paralichthys
dentatus (Summer Flounder), young-of-the-year (YOY) Pomatomus saltatrix (Bluefish),
Prionotus evolans (Striped Searobin), Stenotomus chrysops (Scup), Scophthalmus aquosus
(Windowpane Flounder), Raja eglanteria (Clearnose Skate), and Morone saxatilis
(Striped Bass). Temporal diet composition was estimated for the consistently abundant
YOY Bluefish, Summer Flounder, and Scup, where most nseason > 25. Subsampling of
large catches of YOY Bluefish and Scup led to investigation of diet composition by
cluster sampling. Important prey included Crangon sp. (sand shrimp), Cancer irroratus
(Rock Crab), and forage fishes. Pseudopleuronectes americanus (Winter Flounder), once
a common prey item in stomachs of piscivorous Long Island fishes, contributed ≤ 6.7 %O
and ≤ 1.6 %W to the diets of Summer Flounder, Striped Searobin, Striped Bass, and YOY
Bluefish. These changes may be due to shifts in the abundance of prey items or changes
in spatial overlap of predator and prey.
Ecosystem-based management of fish stocks using multispecies models has
gained favor in recent decades. This approach yields information about sustainability
while incorporating the effects of ecological processes such as predation
among interacting populations (Latour et al. 2003, Link 2002). The complexity
of multispecies models poses several challenges to managers and scientists,
including greater parameterization and additional data requirements (Latour
et al. 2003). A quantitative understanding of piscivorous predation on fishes is
paramount in understanding trophic dynamics and constructing food-web models
utilized by ecosystem-based management (Pauly et al. 2000, Steimle et al. 2000).
The State of New York has recently invested significant resources into developing
ecosystem models and infrastructure to support development of ecosystem
management for Long Island’s bays and estuaries (New York Ocean and Great
Lakes Ecosystem Conservation Act 2006). Presently, there is a need to conduct
trophic analyses in support of these activities and to report indices in a consistent
manner so that estimates of predator and prey relationships are comparable both
spatially and temporally (Cortés 1997).
1School of Marine and Atmospheric Sciences, Stony Brook University, Stony Brook, NY
11794. *Corresponding author - email@example.com.
292 Northeastern Naturalist Vol. 18, No. 3
Trophic interactions and feeding habits of fishes have been described in detail
for most mid-Atlantic estuaries and bays, with the exception of Long Island, NY,
waters. With more than 30 bays and 450 km of coastline surrounding Long Island
(Engers and Buckner 2000), trophic interactions among fishes have received far
less attention than in the neighboring Hudson-Raritan estuary of New York and
New Jersey (Buckel and Conover 1997; Buckel et al. 1999; Gardinier and Hoff
1982; Hurst and Conover 2001; Juanes et al. 1993, 1994; Mancini and Able 2005;
Stehlik and Meise 2000; Steimle et al. 2000). The few studies that have examined
feeding habits of predatory fishes in Long Island bays over the last 50 years focused
solely on species collected from Great South Bay, including Paralichthys
dentatus L. (Summer Flounder; Poole 1964), young-of-the-year (YOY) Pomatomus
saltatrix L. (Bluefish; Juanes and Conover 1995, Juanes et al. 1994), and
Prionotus evolans L. (Striped Searobin; Schreiber 1973) or from western Long
Island bays (Manhasset, Jamaica, and Little Neck bays) including Morone saxatilis
Walbaum (Striped Bass) and YOY Bluefish (Buckel and McKown 2002).
A lack of data concerning trophic interactions of fishes in Long Island systems
has hampered a shift towards ecosystem-based fisheries management in the State
of New York as mandated by the New York Ocean and Great Lakes Ecosystem
Conservation Act (ECL Article 14). This recent legislation “encourages scientific
research and information sharing that will inform ecosystem-based management
decisions and enhance ecosystem management capabilities” in New York’s
Coastal Waters (New York Ocean and Great Lakes Ecosystem Conservation Act
2006). The purpose of this paper is to provide an understanding of general and
temporal diet composition of common fishes present in Long Island waters. General
diet composition is reported in standard indices and is examined assuming
simple random sampling, while diet composition via cluster sampling provides
population-level diet composition when appropriate. The presented diet composition
is discussed relative to intermittent historical studies to elucidate long-term
change in the trophic dynamics of Long Island bays.
Materials and Methods
Specimens were collected from three bodies of water around Long Island, NY:
Port Jefferson Harbor (PJH), Great South Bay (GSB), and Shinnecock Bay (SB)
(Fig. 1). PJH, located on the rocky north shore, contains many deep, cool channels
and connects to Long Island Sound through a narrow inlet. It covers an area of
approximately 4 km2, has an average depth of 4.4 m, and experienced an average
salinity of 26 ppt during our 2007 trawl survey (Gross et al. 1972, Sagarese 2009,
USFWS 1997). In contrast, GSB and SB, both situated on the sandy south shore,
are shallow barrier beach and lagoonal estuaries with abundant salt marshes and
tidal flats and direct connections to the Atlantic Ocean through inlets. GSB is
the largest saltwater bay in New York State, covering an area of 235 km2 with
an average depth of 1.3 m (Hinga 2005, Wilson et al. 1991). GSB is influenced
by heavy riverine and groundwater flow and is characterized by salinity ranging
2011 S.R. Sagarese, R.M. Cerrato, and M.G. Frisk 293
from 25–30 ppt (Hinga 2005). SB covers an area of 39 km2, averages less than
2 m in depth, and has an average salinity of 30 ppt (Buonaiuto and Bokuniewicz
2008, Green and Chambers 2007). These three bays were divided into two categories
for statistical analyses due to differences in geologic and oceanographic
conditions: (1) north shore (PJH) and (2) south shore (GSB and SB).
Collection of specimens
Otter trawls and beach seines were employed in PJH and SB to obtain fish for
dietary analysis bimonthly between April and September of 2007 and monthly
between May and September of 2008. For GSB, fishes were collected by otter
trawl in May, July/August, and October of 2007 as part of a fishery independent
survey commissioned by the New York State Department of Environmental Conservation
(Frisk and Munch 2008). A 9-m otter trawl (0.6-cm cod-end) towed
at 2.5 knots for 8 minutes (12 minutes in GSB) per station collected specimens
from the deeper areas and navigation channels of each bay. Trawl stations were
randomly selected by dividing each body of water into numbered boxes of equal
size and using a random-number generator to determine which box would be
sampled. A 61-m beach seine (0.6-cm mesh) collected fishes from intertidal regions
of PJH and SB. Beach seine stations were randomly selected from 500-m
intervals along the perimeter of each site. Active sampling gears were chosen as
they provide more accurate estimates of food consumption by sampling both lowactivity,
“nonforaging” fishes and actively feeding fishes (Cortés 1997). Due to
the failure of the listed gears to capture Striped Bass and the importance of this
predator within Long Island bays, hook-and-line fishing was employed in year
two of this study.
Figure 1. Location of study sites where predatory fishes were collected from Long Island
waters in 2007 and 2008. White indicates land, gray indicates water. PJH = Port Jefferson
Harbor, GSB = Great South Bay, SB = Shinnecock Bay.
294 Northeastern Naturalist Vol. 18, No. 3
Simple random sampling. Diets of fishes captured throughout the sampling
seasons of 2007–2008 were evaluated under the assumption that each fish in the
population was an independent sampling unit with equal probability of capture in
our sampling regime (Cochran 1977). Upon capture, specimens were measured
for total length (TL, to the nearest mm), immediately put on ice, returned to the
laboratory, and frozen as soon as possible. Fishes were later thawed and weighed
(to the nearest g), and their stomachs were extracted and weighed (to the nearest
mg). To determine the stomach content weight for each specimen, extracted
stomachs were weighed both before and after emptying their contents. Intensity
of feeding for each predatory fish was measured via a stomach fullness index
(SFI) (Hureau 1969):
Sfi= (stomach content weight [g] / fish weight [g]) x 100.
Sfivalues were calculated for all specimens regardless of the presence or
absence of stomach contents, to provide unbiased estimates of feeding intensity.
Stomach contents were sorted, and items were identified to the lowest taxa
possible, enumerated, and weighed (to the nearest mg). For each prey item,
standard indices were reported, including percent by number (%N), percent by
weight (%W), and percent frequency of occurrence (%O), with seasonal values
calculated when feasible (Chipps and Garvey 2007, Hyslop 1980). In addition,
an index of relative importance (IRI; Pinkas et al. 1971) was calculated for each
prey item and converted into a percentage to allow for comparisons to other
studies (Cortés 1997). For Stenotomus chrysops L. (Scup), only %W and %O
are reported because their dentition, which consists of hard plates for crushing
prey, makes enumerating individual prey items difficult. The prey category “marine
plant matter” (MPM) consisted of unidentifiable algae and seaweed, while
“terrestrial plant matter” (TPM) included tree branches and other fragments of
terrestrial plants. The “nonliving matter” included plastic debris and rocks, while
the “unknown” grouping consisted of unrecognizable organic matter.
Cumulative prey curves were calculated for all species to determine whether
an adequate number of specimens had been collected to describe the diet (Cortés
1997, Gelsleichter et al. 1999). The order in which stomach contents were
analyzed was randomized ten times, and the mean number of new prey species
(± standard error) was plotted against the cumulative number of fish examined
(Gelsleichter et al. 1999). Graphically, sample size was deemed adequate once an
asymptotic relationship was displayed.
Cluster sampling estimation. Large catches of fishes were subsampled
when necessary and processed for diet composition utilizing cluster-sampling
estimation (Buckel et al. 1999). Cluster sampling presents population-level
diet data by factoring in the relative abundance of species during each sampling
event and taking into account similarities in diet composition among
individual fish (Bogstad et al. 1995, Buckel et al. 1999). We defined a “cluster”
(Cochran 1977) as the diet composition of a fish species captured by tow
(seine or trawl) and assumed each cluster represented an individual sampling
2011 S.R. Sagarese, R.M. Cerrato, and M.G. Frisk 295
unit. For each cluster, an estimate of the mean proportion of stomachs containing
prey type k (Fk; specimens with empty stomachs were neglected in this
calculation), an estimate of the mean proportional contribution of prey type
k by weight (Wk), and estimators of variance were calculated as in Buckel et
al. (1999). An intracluster correlation coefficient (ICC; ρ) was calculated for
each prey item encountered in cluster sampling estimation. This parameter
ranges from 0 to 1 and reflects the relationship of the variance within clusters
w) to the variance between clusters (s2
b) as described in Steel et al. (1997).
An ICC value of 0 is indicative of diverse stomach contents for fish within a
cluster, resulting in greater variance within tows. In contrast, an ICC value of
1 occurs when stomach contents of fish within a cluster are identical, resulting
in greater variance between tows. ICC values close to zero indicate simple random
sampling is an appropriate measure of diet composition whereas as ICC
approaches 1, cluster sampling estimation is necessary to accurately describe
the diet due to the similarity of diet composition within tows.
The diet compositions of fishes were compared between the north and south
shores of Long Island to determine whether important prey items differed between
the two contrasting environments. The non-normal and heterogeneous
nature of our data prevented parametric statistical testing despite transformation
efforts. Instead, a one-way, non-parametric, multivariate analysis of covariance
(NP-MANCOVA) for an unbalanced ANOVA design using Bray Curtis distances
on the weights (in g) of prey items was used to test for regional differences in
diet composition, using the program DISTLM v.5 (Anderson 2001, Braccini et
al. 2005). DISTLM tests for the multivariate null hypothesis of no relationship
between a data matrix and a design matrix that codes for a particular term in
the model and is achieved through multivariate regression analysis on a matrix
of distance measures (McArdle and Anderson 2001, McKinnon et al. 2008).
The test statistic is a multivariate analogue to Fisher's F-ratio and is calculated
directly from a symmetric distance or dissimilarity matrix with the P-value obtained
using permutations (Anderson 2001). For comparisons between sites for
individual species, site was treated as a fixed factor, with the weights of each
prey item as dependent variables and individual fish as replicates. Individual
fish were used as replicates rather than clusters because cluster sampling estimation
was inappropriate for Summer Flounder and Scopthalmus aquosus
Mitchill (Windowpane Flounder). Fish length was chosen as a covariate to
control for its effect on the dependent variables. Significance was determined
by 9999 permutations of the raw data. A significance level of α = 0.05 was set
a priori and used to test for differences in diet composition. We also performed
a NP-MANOVA to test for differences in diet composition between species. For
this analysis, species was treated as a fixed factor and coded as a design matrix
within an input file (Anderson 2001). Individual weights of each prey item were
the dependent variables, while individual fish represented replicates.
296 Northeastern Naturalist Vol. 18, No. 3
Collection of specimens
Between April and October of 2007 and May and September of 2008, a total of
459 otter trawls and 260 beach seines were conducted during daylight throughout
the sites (Table 1, Fig. 2). Fishes were collected over a total of 53 days: 21 in PJH,
10 in GSB, and 22 in SB. YOY Bluefish, Summer Flounder, Scup, Windowpane
Flounder, Striped Searobin, Striped Bass, and Raja eglanteria Bosc (Clearnose
Skate) were chosen for dietary analyses (Table 2). The majority of specimens
were collected by otter trawl (97%), except for YOY Bluefish and Striped Bass.
Overall, 84% of YOY Bluefish were collected in beach seines. While 22% of
Striped Bass were collected in beach seines from PJH in 2008, additional samples
(n = 18) were obtained by hook and line from other south shore locations, including
Moriches Bay (n = 14) and Montauk Point (n = 4).
Simple random sampling. Gut contents of 523 fishes encompassing seven
species were examined to determine general diet patterns over this two-year
study (Table 2). Striped Bass had the highest proportion of empty stomachs
(35%), while Clearnose Skate had the lowest (10%). Of the seven species
examined, Summer Flounder exhibited the highest average Sfi(5.64 ± 3.91,
S.E.) followed by Scup (4.67 ± 0.64, S.E.), while Windowpane Flounder had
the lowest average Sfi(0.71 ± 0.11, S.E.). Overall, 58 types of prey were
identified, including 24 species of teleosts and 13 species of crustaceans (Supplemental
appendices 1 and 2, available online at http://www.eaglehill.us/
NENAonline/suppl-files/n18-3-972-Sagarese-s1, and, for BioOne subscribers,
at http://dx.doi.org/10.1656/N972.s1). Important teleost prey
included Menidia menidia L. (Atlantic Silverside), Brevoortia tyrannus
Table 1. Summary of effort and gear used to collect common fishes for dietary analysis from Long
Island waters in 2007 and 2008. See Fig. 1 for locations of individual sites. PJH = Port Jefferson
Harbor, GSB = Great South Bay, SB = Shinnecock Bay.
Year Gear Sites sampled Number of nets set Months sampled Depth range (m)
2007 Otter trawl PJH 118 Apr–Sep 2.1–20.0
2007 Otter trawl GSB 99 May–Oct 1.6–12.9
2007 Otter trawl SB 79 Apr–Aug 1.6–13.6
2007 Beach seine PJH 89 May–Sep 0.0–3.0
2007 Beach seine SB 81 Apr–Aug 0.0–3.0
2008 Otter trawl PJH 62 May–Sep 3.2–23.0
2008 Otter trawl SB 101 May–Sep –
2008 Beach seine PJH 48 Jun–Sep 0.0–3.0
2008 Beach seine SB 42 Jun–Sep 0.0–3.0
Figure 2 (opposite page). Locations where predatory fishes were captured by otter trawl
or beach seine by site in 2007 and 2008. Triangles represents otter trawls, circles indicate
beach seines, open symbols indicate 2007, and solid symbols represent 2008. White indicates
land, gray indicates water. A) PJH, B) GSB, and C) SB.
2011 S.R. Sagarese, R.M. Cerrato, and M.G. Frisk 297
298 Northeastern Naturalist Vol. 18, No. 3
Latrobe (Atlantic Menhaden), and Anchoa mitchilli Valenciennes (Bay Anchovy)
(Table 3; Supplemental Appendix 1, available online at http://
www.eaglehill.us/NENAonline/suppl-files/n18-3-972-Sagarese-s1, and, for
BioOne subscribers, at http://dx.doi.org/10.1656/N972.s1). Although all
species fed on teleosts in varying amounts, with YOY Bluefish consuming the
highest proportion, crustaceans dominated the diets of most fishes (Fig. 3).
Summer Flounder and Striped Searobin preyed heavily on Crangon sp. (sand
shrimp), while Neomysis sp. (mysid shrimp) dominated Windowpane Flounder
diet (Table 3). Cancer irroratus Say (Rock Crab) was important in the diet
of Clearnose Skate (Table 3). Striped Bass foraged predominately on teleosts
including Summer Flounder and Sphoeroides maculatus Bloch and Schneider
(Northern Puffer) (Table 3; Supplemental appendices 1 and 2, available
online at http://www.eaglehill.us/NENAonline/suppl-files/n18-3-972-Sagarese-
s1, and, for BioOne subscribers, at http://dx.doi.org/10.1656/N972.
s1). Clams, particularly of the genus Gemma, overwhelmed the Scup diet both
by weight and frequency (Table 3). For most species, cumulative prey curves
exhibited an upward trend, indicating that the number of stomachs analyzed
may not fully describe overall trophic diversity (Fig. 4).
The diets of Scup, YOY Bluefish, and Summer Flounder were examined using
%W and %O by season (Fig. 5). For 2007, seasons were defined as follows:
spring (1 May to 20 June), summer (21 June to 22 September), and fall (23
September to 31 October). For 2008, seasons differed slightly as follows: spring
(1 May to 19 June), summer (20 June to 21 September), and fall (22 September
to 31 October). Small overall sample sizes (<50) coupled with sporadic catches
over time prevented temporal analysis on the remaining 4 species.
Table 2. Descriptive statistics of common fishes examined for dietary analysis. n = sample size,
Empty = number of empty stomachs, TL range = size range of specimens, TLavg = average total
length, Wavg = average weight of fish, Savg = average stomach content weight, SFIavg = average stomach
fullness index, ± indicates standard error.
Species n Empty (%) (cm) TLavg (cm) Wavg (kg) Savg (g) SFIavg
Bluefish 191 23 (12%) 12.0–23.0 17.64 0.06 1.56 2.28
± 0.16 ± 0.00 ± 0.17 ± 0.18
Summer Flounder 141 29 (21%) 26.0–64.9 38.22 0.68 5.55 5.64
± 0.61 ± 0.03 ± 1.04 ± 3.91
Scup 70 11 (16%) 17.5–37.0 25.54 0.32 1.42 4.67
± 0.54 ± 0.02 ± 0.22 ± 0.64
Windowpane Flounder 42 6 (14%) 21.0–31.6 26.60 0.26 1.77 0.71
± 0.43 ± 0.01 ± 0.30 ± 0.11
Striped Searobin 36 4 (11%) 20.7–42.6 31.44 0.45 5.08 0.92
± 0.84 ± 0.03 ± 1.16 ± 0.17
Striped Bass 23 8 (35%) 39.6–95.9 68.56 3.52 24.79 0.90
± 4.67 ± 0.58 ± 10.65 ± 0.28
Clearnose Skate 20 2 (10%) 49.5–71.0 63.34 1.58 20.74 1.31
± 0.07 ± 1.10 ± 2.40 ± 0.15
2011 S.R. Sagarese, R.M. Cerrato, and M.G. Frisk 299
Most Scup were collected during spring (n = 42), with fewer individuals obtained
in summer (n = 28). No collections were made in 2008. Scup diet varied
as mollusks, particularly clams of the genus Gemma, dominated weight and
frequency during spring and summer, with crustaceans increasing in importance
during summer (Fig. 5a, b).
YOY Bluefish collections were divided into 3 seasons: summer 2007 (n =
46), fall 2007 (n = 84), and summer 2008 (n = 61). Collections increased during
summer for both years as they recruited to our fishing gear. Teleost prey was
the largest diet component for all seasons examined (Fig. 5c, d). YOY Bluefish
collected in summer 2007 consumed mainly Bay Anchovy, while those collected
in fall 2007 preyed heavily on Atlantic Menhaden and Silverside. YOY Bluefish
collected in summer 2008 consumed modest amounts of Cynoscion regalis Bloch
and Schneider (Weakfish) and Atlantic Silverside. During summer 2008, crustaceans
peaked in importance in YOY Bluefish diets as sand shrimp consumption
increased (Fig. 5c, d).
Summer Flounder collections were divided into 6 seasons with highly variable
sample sizes: spring 2007 (n = 9), summer 2007 (n = 55), fall 2007 (n = 4),
spring 2008 (n = 19), summer 2008 (n = 52), and fall 2008 (n = 2). The Summer
Flounder diet was generally focused on crustaceans and teleosts over all time
periods (Fig. 5e, f). Sand shrimp was common during most months, while other
crustaceans were patchy in occurrence. Lysiosquilla sp. (mantis shrimp) consumption
increased during summer 2008. Multiple teleosts contributed to diet
composition in each season. Ammodytes sp. (sand lance) was important in diets
Figure 3. Comparison of major prey types given in percent index of relative importance
(%IRI) for each predatory fish. Scup is excluded from this analysis due to the difficulty
in obtaining %N estimates.
300 Northeastern Naturalist Vol. 18, No. 3
Table 3. Abbreviated diet composition of fishes in Long Island bays expressed as percent by number (%N), percent by weight (%W), percent frequency of
occurrence (%O), and percent index of relative importance (%IRI). UNID = unidentified, MPM = marine plant matter, TPM = terrestrial plant matter.
Striped Bass (n = 23) Bluefish (n = 191) Striped Searobin (n = 36)
Common name %N %W %O %IRI %N %W %O %IRI %N %W %O %IRI
Bay Anchovy, Anchoa mitchilli (Valenciennes) 0.00 0.00 0.00 0.00 8.78 12.97 8.33 3.76 0.00 0.00 0.00 0.00
Atlantic Menhaden, Brevoortia tyrannus (Latrobe) 0.00 0.00 0.00 0.00 19.95 31.55 26.79 28.65 0.00 0.00 0.00 0.00
Weakfish, Cynoscion regalis (Bloch and Schneider) 0.00 0.00 0.00 0.00 1.86 7.54 4.17 0.81 0.00 0.00 0.00 0.00
Atlantic Silverside, Menidia menidia (L.) 0.65 0.02 6.67 0.09 34.04 33.04 33.93 47.27 0.17 0.64 3.13 0.02
Summer Flounder, Paralichthys dentatus (L.) 2.58 45.12 20.00 20.05 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00
Winter Flounder, Pseudopleuronectes americanus (Walbaum) 0.65 0.30 6.67 0.13 0.27 0.73 0.60 0.01 0.17 0.41 3.13 0.02
Northern Pipefish, Syngnathus fuscus (Storer) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.17 0.22 3.13 0.01
Other teleosts 7.10 32.54 33.33 27.77 1.60 1.97 3.57 0.26 1.34 2.88 6.25 0.22
UNID Fish 5.16 3.21 33.33 5.87 16.49 8.63 30.95 16.15 0.33 1.03 3.13 0.04
Sand shrimp, Crangon sp. 52.90 3.37 13.33 15.77 5.59 2.68 10.12 1.74 69.57 70.42 81.25 95.58
Mysid shrimp, Neomysis sp. 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 22.91 0.54 3.13 0.62
Rock Crab, Cancer irroratus (Say) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 1.84 1.33 9.38 0.25
Spider Crab, Libinia emarginata (Leach) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00
Other crustaceans 20.65 10.61 26.67 17.52 2.39 0.21 0.00 0.00 1.17 21.78 15.63 3.01
Annelids 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00
Blue Mussel, Mytilus edulis (L.) 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00
Razor Clam, Ensis directus (Conrad) 1.94 0.59 13.33 0.71 0.00 0.00 0.00 0.00 0.67 0.05 6.25 0.04
Other mollusks 1.29 0.42 13.33 0.48 0.00 0.00 0.00 0.00 0.17 0.12 3.13 0.01
Other prey categories
Eel Grass, Vallisneria americana Michx. 0.00 0.00 0.00 0.00 1.86 0.30 4.17 0.19 0.17 0.01 3.13 0.00
MPM 0.65 0.00 6.67 0.09 4.26 0.17 9.52 0.88 0.17 0.02 3.13 0.00
TPM 0.00 0.00 0.00 0.00 0.27 0.02 0.60 0.00 0.17 0.20 3.13 0.01
Nonliving matter 6.45 3.82 53.33 11.52 0.53 0.01 1.19 0.01 1.00 0.34 15.63 0.18
Unknown 0.00 0.00 0.00 0.00 2.13 0.17 5.36 0.26 0.00 0.00 0.00 0.00
Other 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00
2011 S.R. Sagarese, R.M. Cerrato, and M.G. Frisk 301
Table 3, continued.
Clearnose Skate (n = 20) Summer Flounder (n = 141) Windowpane Flounder (n = 42) Scup (n = 70)
Common name %N %W %O %IRI %N %W %O %IRI %N %W %O %IRI %W %O
Bay Anchovy 0.00 0.00 0.00 0.00 4.81 2.30 4.46 1.11 0.02 0.23 2.70 0.01 0.00 0.00
Atlantic Menhaden 0.00 0.00 0.00 0.00 0.94 1.09 1.79 0.13 0.00 0.00 0.00 0.00 0.96 6.78
Weakfish 0.00 0.00 0.00 0.00 0.42 2.98 4.46 0.53 0.00 0.00 0.00 0.00 0.00 0.00
Altantic Silverside 0.30 0.32 5.56 0.02 2.93 3.94 7.14 1.71 0.00 0.00 0.00 0.00 0.00 0.00
Summer Flounder 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00
Winter Flounder 0.00 0.00 0.00 0.00 0.52 1.60 4.46 0.33 0.00 0.00 0.00 0.00 0.00 0.00
Northern Pipefish 0.00 0.00 0.00 0.00 0.21 0.08 1.79 0.02 0.02 0.08 2.70 0.00 0.00 0.00
Other Teleost 0.30 0.68 5.56 0.04 5.23 16.41 16.07 12.14 0.00 0.00 0.00 0.00 0.00 0.00
UNID Fish 0.89 0.23 16.67 0.13 3.45 12.36 16.96 9.36 0.02 0.01 2.70 0.00 2.09 1.69
Sand Shrimp 42.01 9.85 72.22 25.84 25.21 12.58 32.14 42.39 2.90 36.09 51.35 14.91 2.46 6.78
Mysid Shrimp 0.00 0.00 0.00 0.00 39.54 1.09 2.68 3.80 96.49 59.90 72.97 85.01 2.64 13.56
Rock Crab 38.17 64.59 94.44 66.95 5.96 13.93 10.71 7.44 0.00 0.00 0.00 0.00 0.13 1.69
Spider Crab 1.78 14.69 27.78 3.16 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00
Other Crustaceans 10.95 9.29 22.22 3.10 3.03 19.48 21.43 16.83 0.22 0.31 8.11 0.03 7.94 13.56
Annelids 0.00 0.00 0.00 0.00 1.57 2.90 7.14 1.11 0.00 0.00 0.00 0.00 10.34 15.25
Blue Mussel 4.44 0.22 22.22 0.71 2.72 0.68 8.93 1.06 0.02 0.00 2.70 0.00 0.00 0.00
Clam 0.00 0.00 0.00 0.00 0.10 0.01 0.89 0.00 0.02 0.02 2.70 0.00 62.30 57.63
Other Mollusks 0.00 0.00 0.00 0.00 0.63 7.74 5.36 1.56 0.00 0.00 0.00 0.00 0.07 1.69
Other prey categories
Eel Grass 0.30 0.04 5.56 0.01 0.73 0.14 6.25 0.19 0.02 0.00 2.70 0.00 0.00 0.00
MPM 0.30 0.05 5.56 0.01 0.63 0.09 5.36 0.13 0.04 0.10 5.41 0.01 0.03 1.69
TPM 0.30 0.03 5.56 0.01 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00
Nonliving matter 0.30 0.01 5.56 0.01 0.21 0.23 1.79 0.03 0.17 1.17 2.70 0.03 0.00 0.00
Unknown 0.00 0.00 0.00 0.00 0.73 0.30 3.57 0.13 0.02 0.05 2.70 0.00 5.55 22.03
Other 0.00 0.00 0.00 0.00 0.42 0.06 3.57 0.06 0.06 2.12 5.41 0.09 5.49 18.64
302 Northeastern Naturalist Vol. 18, No. 3
collected during spring in both years, Scup was important during summer 2008,
Bay Anchovy was mainly consumed during spring and summer 2008, Weakfish
dominated diets during summer in both years, and Atlantic Silverside was preyed
upon in small amounts for most sampling seasons.
Cluster sampling estimation. For YOY Bluefish, examination of 26 clusters revealed
Atlantic Silverside and Menhaden dominated mean proportion of frequency
Figure 4. Randomized cumulative prey curves for a) YOY Bluefish , b) Summer Flounder,
c) Scup, d) Windowpane Flounder, e) Striped Searobin, f) Striped Bass and g) Clearnose
Skate. Mean values are plotted, error bars represent ± SE.
2011 S.R. Sagarese, R.M. Cerrato, and M.G. Frisk 303
and mean proportion by weight (Fig. 6a). ICC values for all 19 prey items were
close to 1, indicating cluster sampling estimation was the preferred method to
describe YOY Bluefish diet. For Scup, examination of 24 clusters shows the importance
of clams within Scup diets by frequency and weight (Fig. 6b). ICC values for
15 prey items ranged from 0.01 (ρClam) to 1.00 (ρMPM), indicating cluster sampling
more accurately reflects the dietary preference of Scup.
Statistical analysis was limited to 3 of the 7 species examined due to inadequate
sample sizes (nsite < 15) for both sampling regions (north vs. south).
Windowpane Flounder, YOY Bluefish, and Summer Flounder were examined for
differences in prey items between sites (Table 4). No significant difference in diet
Figure 5. Percent by weight (%W) and percent frequency of occurrence (%O) of potential
prey contributing to the diets of common fishes in Long Island waters by season. Scup:
(a) %W, (b) %O; YOY Bluefish: (c) %W, (d) %O; and Summer Flounder (e) %W, (f) %O.
SP = spring, SU = summer, and FA = fall, 07 = 2007, 08 = 2008.
304 Northeastern Naturalist Vol. 18, No. 3
2011 S.R. Sagarese, R.M. Cerrato, and M.G. Frisk 305
composition between sites was detected for Windowpane Flounder (P = 0.0694).
In contrast, a significant difference was detected in Bluefish (P = 0.0002) and
Summer Flounder (P = 0.0043). Upon examination, YOY Bluefish in north shore
waters consumed a greater mass of sand shrimp and Atlantic Silverside but less
Bay Anchovy. Summer Flounder in north shore waters consumed a greater mass of
Atlantic Silverside, sand lance, and squid. In addition, due to the large difference
in sample size between sites, Summer Flounder diet from the south was much
more diverse with 18 additional prey items identified. The NP-MANOVA test for
differences in the weights of prey items among all seven species pooled over sites
revealed a significant difference between the diet compositions of all predators
(P = 0.0001).
This study analyzed the trophic habits of important finfish for the inshore bays
of Long Island, providing critical data for the development of ecosystem-based
management for an understudied system. Cumulative prey curves indicated that
the number of stomachs examined was sufficient to identify important prey items;
however, for some species, results of this study should be interpreted with caution
due to small overall sample sizes, which made quantifying overall trophic
diversity unfeasible. Diet composition differed significantly between species,
with YOY Bluefish and Striped Bass consuming teleosts, Scup consuming bivalves,
and the remaining four fish species feeding heavily on crustaceans. The
energetic demands of upper trophic finfish occurring in Long Island bays are met
by crustaceans and, to a lesser extent, forage fishes such as Atlantic Silverside
Figure 6 (opposite page). Mean proportion of stomachs containing prey type k (Fk) and
mean proportional contribution of a prey type by weight (Wk) in the diets of common
fishes collected from Long Island waters in 2007 and 2008. A) YOY Bluefish, B) Scup.
nc = total number of clusters collected, nt = total number of specimens. Standard error
estimates, represented by error bars, were calculated from cluster sampling variance estimates,
and all were less than 0.06%. UNID = unidentified, MPM = marine plant matter,
TPM = terrestrial plant matter, NLM = nonliving matter.
Table 4. NP-MANCOVA testing for the effects of site (north versus south) on the weight of prey
items in the diet of predatory fishes while removing the effect of fish length. Site is treated as a
fixed factor, the weights of each prey item are the dependent variables, individual fish represent
replicates, and fish length is used as a covariate. df = degrees of freedom, SS = sums of squares,
P = probability, significance level of α = 0.05 set a priori.
Sample size by site Permutation Monte
Species North South Total df Total SS Pseudo-F P Carlo P
Windowpane Flounder 19 23 41 149488.3 1.9569 0.0694 0.0749
Bluefish 97 94 190 824514.2 4.2334 0.0002 0.0001
Summer Flounder 41 100 140 639663.1 2.5021 0.0043 0.0048
306 Northeastern Naturalist Vol. 18, No. 3
and Menhaden (Nuttall 2010). Sand shrimp is a key prey item within this system,
as it was preyed upon by all seven species examined and was critical in the diets
of Summer Flounder, Striped Searobin, Windowpane Flounder, and Clearnose
Skate. The trophic importance of sand shrimp has been recognized in other New
York ecosystems including the Hudson-Raritan estuary (Steimle and Terranova
1991, Steimle et al. 2000) and previous studies conducted in GSB (Juanes and
Conover 1995, Juanes et al. 2001, Poole 1964). Qualitative comparisons to historical
data collected in the 1950s and 1980s indicated a shift in Summer Flounder
and YOY Bluefish diet from the once-abundant Pseudopleuronectes americanus
Walbaum (Winter Flounder) to more common forage fishes and crustaceans.
Historically, YOY Winter Flounder has been an important prey item in GSB
(Juanes and Conover 1995, Poole 1964, Schreiber 1973). However, our study
indicated only 1.5% of the total number of predator stomachs examined contained
a single YOY Winter Flounder, making them a rare prey item for this
area. Although many factors can complicate comparisons between studies, such
as different predator size classes or sample sizes, our results are consistent with
a drastic decline of Winter Flounder observed in the bays of Long Island by annual
surveys conducted by NYSDEC (Socrates 2006). Our data indicated that
predators are consuming Winter Flounder in far less amounts than previously
observed in terms of %W and %O. Dietary analysis of summer-caught Summer
Flounder (n = 1210, 24–68 cm TL) in 1958 and 1959 identified Winter Flounder
as an important prey item in GSB (27.8%W, 7.4%O) (Poole 1964). Examination
of Summer Flounder stomachs captured from the south shore (n = 97, 26–65 cm
TL) revealed a decline in consumption of Winter Flounder (2.4%W, 5.2%O).
Winter Flounder was also more important to the diet of YOY Bluefish in 1989
(11.1%W) (Juanes and Conover 1995) compared with 2007–2008 (1.5%W).
This study indicated that adult Summer Flounder (>28 cm TL; O’Brien et
al. 1993) in Long Island waters exhibit a highly diverse diet as evidenced by the
presence of 14 teleost and 9 arthropod species. This species primarily consumed
crustaceans, notably sand shrimp, mantis shrimp, and Rock Crab. Our analysis
differed from the species adult diet composition (n = 88, TLavg = 48 cm) for
offshore waters south of New England (Bowman et al. 2000). Offshore, sand
lance and squid are important prey items, while crustaceans are rarely consumed,
possibly a result of differences in predator size or due to the nature of the ocean
environment (Bowman et al. 2000). Our findings also contrast with an inshore
study conducted by Poole (1964), who observed summer caught adult Summer
Flounder feeding heavily on teleosts, including Winter Flounder and Syngnathus
fuscus Storer (Northern Pipefish) in GSB between 1958–1959. As Winter
Flounder have recently declined in Long Island waters (Frisk and Munch 2008,
Socrates 2006), this difference in teleost consumption may reflect a recent shift
in diet to more abundant, smaller forage fishes. Since Summer Flounder tend to
prefer demersal teleost prey (Manderson et al. 2000), fewer encounters with demersal
species such as Winter Flounder may force this predator to consume other
benthic prey or pelagic fishes. Summer Flounder exhibited the highest Sfiand
commonly consumed teleosts and crustaceans over a range of sizes.
2011 S.R. Sagarese, R.M. Cerrato, and M.G. Frisk 307
Poole (1964) acknowledged the importance of sand and mysid shrimp in GSB
during the 1950s. Our results for Summer Flounder provide supporting evidence
that sand shrimp is an important prey item in GSB. Sand shrimp is a principal
link in the nutrient cycle of many coastal ecosystems and its high abundance and
growth rates during warm summer months make them ideal prey (Latour et al.
2008, Modlin 1980, Steimle et al. 2000, Wilcox and Jeffries 1973). Similar to
Latour et al. (2008), who analyzed Summer Flounder diet (n = 3079, 12–73 cm TL)
in Chesapeake Bay, we observed high sand shrimp predation in spring and summer.
However, in our study, Summer Flounder diet shifted to other crustaceans, including
Rock Crab and mantis shrimp, during late summer/fall, reflecting a possible
decline in sand shrimp abundance. The difference we found in diet composition between
north- and south-shore Summer Flounder may reflect differences in regional
abundances or spatial overlap between predator and prey.
The examination of YOY Bluefish from Long Island waters proved different
from other predators as this species mainly consumed teleosts, including Atlantic
Silverside and Menhaden, and occasionally fed upon crustaceans. Bluefish and
Atlantic Silversides are both present inshore in similar habitat during early spring
and remain abundant during summer, making these forage fish available to YOY
Bluefish (Juanes and Conover 1995). Our results agree with previous findings for
New York (Buckel and McKown 2002, Juanes and Conover 1995, Juanes et al.
2001) and New Jersey waters (Scharf et al. 2004). One unique observation from
our study was that sand shrimp were rare in YOY Bluefish stomach contents with
the exception of summer 2008. In contrast, other studies found sand shrimp importance
to rival teleost in the late 1980s in GSB (Juanes and Conover 1995) and
the Sandy Hook Bay estuary of New Jersey (Friedland et al. 1988). Consumption
of shrimp depends on their relative abundance and size relative to fish prey
(Juanes et al. 2001). Laboratory studies showed that YOY Bluefish selectively
ingested fish compared with similarly-sized shrimp prey, which required added
manipulation and longer handling times (Juanes et al. 2001). Ingesting teleost
prey results in higher feeding and growth rates and improves YOY Bluefish body
condition (Buckel et al. 1999, Friedland et al. 1988, Juanes and Conover 1994).
Adult Striped Searobins (>22.1 cm fork length [FL], McEachran and Davis
1970) were collected between May and July, and therefore our results reflect a
spring/early summer diet. Striped Searobin displayed the most consistent diet
by feeding almost exclusively on sand shrimp and rarely consuming teleosts.
Partially buried sand shrimp are extremely vulnerable to bottom-feeding searobins,
which use modified pectoral fins to search and flush out prey items from
the sediment (Manderson et al. 1999). A previous Long Island study reported
a more diverse diet for Striped Searobin (n = 40) from GSB, with sand shrimp
and Atlantic Silverside common in 1972 (Schreiber 1973). However, this difference
may be attributed to collection of specimens at different times of the year,
as Schreiber (1973) does not report month of capture. A conflicting result was
observed in Long Island Sound by Richards et al. (1979), who collected Striped
Searobins (n = 390, 12–33 cm TL) over irregular intervals during 1971–1973 and
1976–1977 and concluded that crabs were more abundant in stomachs than sand
308 Northeastern Naturalist Vol. 18, No. 3
shrimp. This difference may result from different hydrographic conditions of
temperature, salinity, and circulation encountered in the deeper Sound compared
to inshore bays or may reflect an ontogenetic shift in diet. The dominance of sand
shrimp in the diet of Striped Searobin agrees with observations from neighboring
estuaries. Within the lower Hudson-Raritan estuary, sand shrimp was the
predominant prey item for specimens (n = 153, 4–47 cm TL) collected in 1996
and 1997 (Steimle et al. 2000). In the Navesink River/Sandy Hook Bay estuarine
system, sand shrimp occurred in 81% of Striped Searobin stomachs (n = 78,
12–37 cm TL; Manderson et al. 1999).
This study is the first documentation of the diet of adult Scup (fl> 15.5 cm;
O’Brien et al. 1993, Steimle et al. 1999) inhabiting inshore waters of Long Island.
Bivalves and annelids were identified as important prey items. Our results differ
from reports of adult Scup diet (n = 111, TLavg = 16 cm) in offshore waters of southern
New England, where animal remains and polychaetes made up a majority of
the diet (Bowman et al. 2000). Between 1981 and 1990, dietary analysis of Scup
(n = 330, 11–40 cm TL) revealed that annelids and amphipods were essential prey
items, with mollusks of occasional importance (Steimle et al. 1999). The predominance
of bivalves in the diet of Scup was also absent from specimens collected
inshore within the lower Hudson-Raritan estuary. Scup (n = 254, 8–24 cm FL) fed
upon smaller prey items including unidentified organic matter and mysid shrimp
between July 1996 and November 1997 (Steimle et al. 2000). However, Steimle
et al. (2000) chose to report Scup diet by pooling results from juveniles (<15.5 cm
FL) and adult Scup (>15.5 cm FL), which may have masked differences between
the diets of different life stages. High bivalve presence in Scup diet from Long Island
bays during early summer followed by a drop off in late summer may parallel
the abundance of these benthic organisms. The temporal trends in clam consumption
may correspond to the abundance of YOY and juvenile clams during periods
of high nutritive value of sediment that promotes benthic growth in early summer
more effectively than later in the season (Cheng et al. 1993).
Here we provide the first description of adult Windowpane Flounder (>21.2
cm TL; O’Brien et al. 1993) diet for bay waters of Long Island. Although the
cumulative prey curve appeared close to an asymptote, a larger sample size
would have enhanced descriptive power. This species primarily consumed
mysid shrimp and to a lesser extent sand shrimp. Our results agree with diet
composition for Windowpane Flounder (n = 132, TLavg = 26 cm) in offshore
waters south of New England where crustaceans including sand and mysid
shrimp were important prey items (Bowman et al. 2000). However, we found
very little teleost consumption, whereas in the offshore environment, sand lance
was classified as an important prey item (Bowman et al. 2000). The dominance
of planktonic mysid shrimp has also been documented in neighboring waters.
Between 1982 and 1985 in the New York Bight, the Windowpane Flounder diet
(n = 131) predominantly consisted of mysid shrimp (Steimle and Terranova
1991). Similarly, Windowpane Flounder (n = 570, 2–35 cm TL) sampled from
the Hudson-Raritan estuary in 1996 and 1997 fed heavily on mysid shrimp,
while also consuming modest amounts of sand shrimp (Steimle et al. 2000).
2011 S.R. Sagarese, R.M. Cerrato, and M.G. Frisk 309
Mysid shrimp cluster near the sea floor during daylight due to their negatively
phototaxic nature and, as a result, are vulnerable to demersal predators like
Windowpane Flounder (Steimle et al. 2000). In addition, Windowpane Flounder
have a small gape size (distance between the maxillary bones in the mouth
interior), which favors the consumption of smaller prey such as mysid shrimp,
as they only reach about 2 cm in length. Windowpane Flounder exhibited the
lowest Sfidue to the consistent consumption of minute crustaceans.
The diet composition of late juvenile and adult (>59 cm TL; Sosebee 2005)
Clearnose Skate reported in this study is the first from Long Island waters. Skates
were collected from Shinnecock Bay between June and August after making their
annual northern migration (Frisk 2010, Sosebee 2005). While a small number of
specimens were obtained, the cumulative prey curve indicates our sample size
was sufficient to identify major prey items. This species primarily consumed
Rock Crab and sand shrimp. Our findings contrast the diet composition for Clearnose
Skate (n = 36, TLavg = 53 cm) in offshore waters of the northwest Atlantic,
where teleosts including Weakfish and flatfishes were important prey items in the
oceanic environment (Bowman et al. 2000). Our results also differ with observations
that crabs (Rock Crab and Ovalipes ocellatus J.F.W. Herbst [Lady Crab])
are more important for Clearnose Skates under 60 cm TL (McEachran 2002).
Seventy percent of our specimens contained Rock Crab in their stomach contents
in varying quantities, possibly reflecting a high abundance of Rock Crab in Long
Island waters. Clearnose Skate diet (n = 96) in Delaware Bay during the summers
of 1989–1994 showed Rock Crab and Ensis directus Conrad (Razor Clam) as
dominant prey items (Steimle et al. 2000). In the lower Hudson-Raritan estuary
during the summers of 1996 and 1997, Clearnose Skate diet (n = 71, 49–86 cm
TL) consisted mostly of Rock Crab and sand shrimp (Steimle et al. 2000). One
similarity between the present study and Steimle et al. (2000) is that most Clearnose
Skate analyzed had contents within their gut as is common for demersal
carnivores (Wetherbee and Cortés 2004).
The last predatory fish species we examined was Striped Bass, but because
of small sample sizes, results are preliminary. Although a small sample size prevented
in-depth examination of spatial and temporal trends, we chose to present
our results because this species is currently an important predator in the system.
Concurrent with an increase in Atlantic Striped Bass from 10 million fish in the
early 1980s to over 50 million fish in recent years (ASMFC 2010, Hartman and
Margraf 2003, Shepard 2006), increased abundance is reflected in seine surveys
conducted by New York State along the south shore of Long Island (Socrates
2006). Striped Bass primarily consumed teleosts including Summer Flounder and
occasionally fed upon sand shrimp. This species exhibited the highest proportion
of empty stomachs likely due to capture method, as passive gear such as hook and
line tends to attract individuals in a population that are actively feeding (Cortés
1997, Wetherbee and Cortés 2004). Steimle et al. (2000) analyzed Striped Bass
diet (n = 280, 13–65 cm TL) between 1996 and 1997 and identified sand shrimp
as the most important prey item, while observing some teleost consumption
throughout the time period.
310 Northeastern Naturalist Vol. 18, No. 3
Throughout our intensive sampling seasons, we were unable to obtain adequate
sample sizes for in-depth study of all fishes chosen for analysis. While our gear
types proved capable of capturing predatory fishes, with the exception of Striped
Bass, additional sampling effort and supplementary gears such as gill nets may
have increased sample sizes. In summary, the general diet composition results
presented in this study are intended to provide a snapshot of the Long Island environment,
showing the role of each species within the food web of Long Island
bays. The observed diet information for predatory fishes has important implications
for ecosystem analysis and management of Long Island bays (Nuttall 2010).
Development of ecosystem models of GSB have relied on data from neighboring
ecosystems, which may result in inaccurate representation of the system’s trophic
dynamics (Nuttall 2010). For example, in the case of Scup, utilization of previous
diet-composition studies would have underrepresented the importance of bivalves
and overrepresented the importance of annelids within GSB. This study provides
diet information specific to the Long Island ecosystem and highlights regional differences
in diet composition for adult Summer Flounder and YOY Bluefish. Our
results for Windowpane Flounder, Clearnose Skate, and adult Scup are the first
reported diet composition within Long Island waters. Continued investigation
of general, temporal, and ontogenetic diet shifts of fishes should be undertaken
to continue building a sufficient diet-composition database for underrepresented
fishes in Long Island bays for use in multispecies models and by managers.
We thank graduate students M. Yencho, M. Nuttall, C. Martinez, and C. Hall, and
many undergraduate students, especially A. Uhlich, for assisting with field sampling.
A. Jordaan provided comments to improve this manuscript. Also, we are grateful for the
hard work of Captains M. Wiggins and D. Bowman from the Marine Science Research
Center and D. Getz, R. McIntyre, and B. Gagliardi from the Stony Brook-Southampton
Marine Station. Funding for this project was provided by the New York State Department
of Environmental Conservation.
Anderson, M.J. 2001. A new method for non-parametric multivariate analysis of variance.
Austral Ecology 26:32–46.
Bogstad, B., M. Pennington, and J.H. Volstad. 1995. Cost-efficient survey designs for
estimating food consumption by fish. Fisheries Research 23:37–46.
Bowman, R.E., C.E. Stillwell, W.L. Michaels, and M.D. Grosslein. 2000. Food of Northwest
Atlantic fishes and two common species of squid. NOAA Technical Memorandum
NMFS–F/NE–155. 138 pp.
Braccini, J.M., B.M. Gillanders, and T.I. Walker. 2005. Sources of variation in the feeding
ecology of the Piked Spurdog (Squalus megalops): Implications for inferring
predator-prey interactions from overall dietary composition. ICES Journal of Marine
Buckel, J.A., and D.O. Conover. 1997. Movements, feeding periods, and daily ration of
piscivorous young-of-the-year Bluefish (Pomatomus saltatrix) in the Hudson River
estuary. Fishery Bulletin 95:665–679.
2011 S.R. Sagarese, R.M. Cerrato, and M.G. Frisk 311
Buckel, J.A., and K.A. McKown. 2002. Competition between juvenile Striped Bass and
Bluefish: Resource partitioning and growth rate. Marine Ecology Progress Series
Buckel, J.A., D.O. Conover, N.D. Steinberg, and K.A. McKown. 1999. Impact of age-0
Bluefish (Pomatomus saltatrix) predation on age-0 fishes in the Hudson River estuary:
Evidence for density-dependent loss of juvenile Striped Bass (Morone saxatilis).
Canadian Journal of Fisheries and Aquatic Sciences 56:275–287.
Buonaiuto, F.S., Jr., and H.J. Bokuniewicz. 2008. Hydrodynamic partitioning of a mixed
energy tidal inlet. Journal of Coastal Research 24(5):1339–1348.
Cheng, I.J., J.S. Levinton, M. McCartney, D. Martinez, and M.J. Weissburg. 1993. A
bioassay approach to seasonal variation in the nutritional value of sediment. Marine
Ecology Progress Series 94:275–285.
Chipps, S.R., and J.E. Garvey. 2007. Assessment of diets and feeding patterns. Pp. 473–
514, In C.S. Guy and M.L. Brown (Eds.). Analysis and Interpretation of Freshwater
Fisheries Data. American Fisheries Society, Bethesda, MD. 961 pp.
Cochran, W.G. 1977. Sampling Techniques, 3rd Edition. John Wiley and Sons, New
York, NY. 428 pp.
Cortés, E. 1997. A critical review of methods of studying fish feeding based on analysis
of stomach contents: Application to elasmobranch fishes. Canadian Journal of Fisheries
and Aquatic Sciences 54:726–738.
Engers, J., and B. Buckner. 2000. The Great Book of Wildfowl Decoys, 1st Edition. Globe
Pequot Press, Guilford, CT. 320 pp.
Friedland, K.D., G.C. Garman, A.J. Bejda, A.L. Studholme, and B. Olla. 1988. Interannual
variation in diet and condition in juvenile Bluefish during estuarine residency.
Transactions of the American Fisheries Society 117:474–479.
Frisk, M.G. 2010. Life-history strategies of batoids. Pp. 283–318, In J.C. Carrier, J.A.
Musick, and M.R. Heithaus (Eds.). Sharks and their Relatives II: Biodiversity, Adaptive
Physiology, and Conservation. CRC Press, Boca Raton, FL. 736 pp.
Frisk, M.G., and S.B. Munch. 2008. Great South Bay fishery independent survey. School
of Marine and Atmospheric Sciences, Stony Brook University, Stony Brook, NY.
Gardinier, M.N., and T.B. Hoff. 1982. Diet of Striped Bass in the Hudson River Estuary.
New York Fish and Game Journal 29(2):152–165.
Gelsleichter, J., J.A. Musick, and S. Nichols. 1999. Food habits of the Smooth Dogfish,
Mustelus canis, Dusky Shark, Carcharhinus obscurus, Atlantic Sharpnose Shark,
Rhizoprionodon terraenovae, and the Sand Tiger, Carcharias taurus, from the northwest
Atlantic Ocean. Environmental Biology of Fishes 54:205–217.
Godwin, C., W. Laney, N. Meserve, S. Meyers, and G. Shepard. 2010. 2010 Review of
the Atlantic States Marine Fisheries Commission Fishery Management Plan for Atlantic
Striped Bass (Morone saxatilis), 2009 Fishing Year. A report prepared by the
Atlantic Striped Bass Plan Review Team, [PROVIDE LOCATION]. 35 pp.
Green, B.S., and R.C. Chambers. 2007. Maternal effects vary between source populations
in the Atlantic Tomcod, Microgadus tomcod. Marine Ecology Progress Series
Gross, M.G., D. Davies, P.M. Lin, and W. Loeffler. 1972. Characteristics and environmental
quality of six north shore bays, Nassau and Suffolk counties, Long Island,
New York. Marine Sciences Research Center, State University of New York, Stony
Brook, NY. Technical Report Series No. 14. 98 pp.
Hartman, K.J., and F.J. Margraf. 2003. US Atlantic coast Striped Bass: Issues with a
recovered population. Fisheries Management and Ecology 10:309–312.
312 Northeastern Naturalist Vol. 18, No. 3
Hinga, K.R. 2005. Water quality and ecology of Great South Bay (Fire Island National
Seashore Science Synthesis Paper). Technical Report NPS/NER/NRTR—2005/019.
National Park Service, Boston, MA.
Hureau, J.C. 1969. Biologie comparee de quelques poissons anarctiques (Nototheniidae).
Bulletin of the Institut Oceanographique Monaco 68:1−44.
Hurst, T.P., and D.O. Conover. 2001. Diet and consumption rates of overwintering YOY
Striped Bass, Morone saxatilis, in the Hudson River. Fisheries Bulletin 99:545–553.
Hyslop, E.J. 1980. Stomach contents analysis: A review of methods and the application.
Journal of Fish Biology 17:411–429.
Juanes, F., and D.O. Conover. 1994. Rapid growth, high feeding rates, and early piscivory
in young-of-the-year Bluefish, Pomatomus saltatrix. Canadian Journal of
Fisheries and Aquatic Science 51:1752–1761.
Juanes, F., and D.O. Conover. 1995. Size-structured piscivory: Advection and the linkage
between predator and prey recruitment in young-of-the-year Bluefish. Marine Ecology
Progress Series 128:287−304.
Juanes, F., J.A. Buckel, and D.O. Conover. 1994. Accelerating the onset of piscivory: Intersection
of predation and prey phonologies. Journal of Fish Biology 45(Supplement
Juanes, F., R.E. Marks, K.A. McKown, and D.O. Conover. 1993. Predation by age-0
Bluefish on age-0 anadromous fishes in the Hudson River estuary. Transactions of the
American Fisheries Society 122:348–356.
Juanes, F., J.A. Buckel, and F.S. Scharf. 2001. Predatory behaviour and selectivity of a
primary piscivore: Comparison of fish and non-fish prey. Marine Ecology Progress
Latour, R.J., M.J. Brush, and C.F. Bonzek. 2003. Toward ecosystem-based fisheries
management: Strategies for multispecies modeling and associated data requirements.
Latour, R.J., J. Gartland, C.F. Bonzek, and R.A. Johnson. 2008. The trophic dynamics
of Summer Flounder (Paralichthys dentatus) in Chesapeake Bay. Fishery Bulletin
Link, J.S. 2002. Ecological considerations in fisheries management: When does it matter?
Mancini, F.T., III, and K.W. Able. 2005. Food habits of young-of-the-year estuarine
fishes in the middle Atlantic bight: A synthesis. Technical report. Institute of Marine
and Coastal Sciences, Rutgers, The State of New Jersey, Tuckerton, NJ. 113 pp.
Manderson, J.P., B.A. Phelan, A.J. Bejda, L.L. Stehlik, and A.W. Stoner. 1999. Predation
by Striped Searobin (Prionotus evolans, Triglidae) on young-of-the-year Winter
Flounder (Pseudopleuronectes americanus, Walbaum): Examining prey size selection
and prey choice using field observations and laboratory experiments. Journal of Experimental
Marine Biology and Ecology 242:211−231.
Manderson, J.P., B.A. Phelan, A.W. Stoner, and J. Hilbert. 2000. Predator-prey relations
between age-1+ Summer Flounder (Paralichthys dentatus, Linnaeus) and age-0
Winter Flounder (Pseudopleuronectes americanus, Walbaum): Predator diets, prey
selection, and effects of sediments and macrophytes. Journal of Experimental Marine
Biology and Ecology 251:17–39.
McArdle, B.H., and M.J. Anderson. 2001. Fitting multivariate models to community
data: A comment on distance-based redundancy analysis. Ecology 82:290–297.
McEachran, J.D. 2002. Skates. Family Rajidae. Pp. 60−75, In B.B. Collette and G. Klein-
MacPhee (Eds.). Bigelow and Schroeder’s Fishes of the Gulf of Maine. 3rd Edition.
Smithsonian Institution Press, Washington, DC. 748 pp.
2011 S.R. Sagarese, R.M. Cerrato, and M.G. Frisk 313
McEachran, J.D., and J. Davis. 1970. Age and growth of the Striped Searobin. Transactions
of the American Fisheries Society 99:343–352.
McKinnon, A.D., S. Duggan, J.H. Carleton, and R. Böttger-Schnack. 2008. Summer
planktonic copepod communities of Australia's North West Cape (Indian Ocean) during
the 1997–1999 El Nino/La Nina. Journal of Plankton Research 30(7):839–855.
Modlin, R.F. 1980. The life cycle and recruitment of the sand shrimp, Crangon septemspinosa,
in the Mystic River estuary, Connecticut. Estuaries and Coasts 3:1−10.
New York Ocean and Great Lakes Ecosystem Conservation Act. 2006. In New York State
Environmental Conservation Law, Article 14.
Nuttall, M.A. 2010. Historical recount of the Great South Bay ecosystem, Long Island,
New York, and a quantitative assessment of the ecosystem structure of Great South Bay
using Ecopath. M.Sc. Thesis. Stony Brook University, Stony Brook, NY. 200 pp.
O’Brien, L., J. Burnett, and R.K. Mayo. 1993. Maturation of nineteen species of finfish
off the Northeast coast of the United States, 1985–1990. NOAA Technical Report
NMFS 113. National Marine Fisheries Service, Woods Hole, MA. 66 pp.
Pauly, D., V. Christensen, and C. Walters. 2000. Ecopath, ecosim, and ecospace as
tools for evaluating ecosystem impacts of fisheries. ICES Journal of Marine Science
Pinkas, L., M.S. Oliphant, and I.L.K. Iverson. 1971. Food habits of Albacore, Bluefin
Tuna, and Bonito in California waters. California Department of Fish and Game Fish
Poole, J.C. 1964. Feeding habits of the Summer Flounder in Great South Bay. New York
Fish and Game Journal 11:28−34.
Richards, S.W., J.M. Mann, and J.A. Walker. 1979. Comparison of spawning seasons,
age, growth rates, and food of two sympatric species of searobins, Prionotus carolinus
and Prionotus evolans, from Long Island Sound. Estuaries 2:255–268.
Sagarese, S.R. 2009. Inshore movements, residency, and abundance of adult Winter
Flounder, Pseudopleuronectes americanus, and piscivorous predation on young-ofthe-
year Winter Flounder within coastal bays of Long Island and an investigation on
the effect of temperature and photoperiod on vertebral band deposition in Little Skate,
Raja erinacea. M.Sc. Thesis. Stony Brook University, Stony Brook, NY. 151 pp.
Scharf, F.S., J.P. Manderson, M.C. Fabrizio, J.P. Pessutti, J.E. Rosendale, R.J. Chant,
and A.J. Bejda. 2004. Seasonal and interannual patterns of distribution and diet of
Bluefish within a Middle Atlantic bight estuary in relation to abiotic and biotic factors.
Estuaries and Coasts 27(3):426–436.
Schreiber, R.A. 1973. The fishes of Great South Bay. M.Sc. Thesis. State University of
Stony Brook, Stony Brook, NY, 199 pp.
Shepard, G. 2006. Status of Fishery Resources off the Northeastern US: Atlantic Striped
Bass. December 2006. Northeast Fisheries Science Center Resource Evaluation and
Assessment Division. Available online at http://www.nefsc.noaa.gov/sos/spsyn/af/
sbass/. Accessed August 2010.
Socrates, J.B. 2006. A study of the Striped Bass in the Marine District of New York
State. New York State Department of Environmental Conservation, East Setauket,
NY. Completion Report for Project AFC-30.
Sosebee, K.A. 2005. Maturity of skates in Northeast United States waters. Journal of
Northwest Atlantic Fishery Science 35:141–153.
Steel, R.G.D., J.H. Torrie, and D.A. Dickey. 1997. Principles and Procedures of Statistics:
A Biometrical Approach. 3rd Edition. McGraw-Hill, New York, NY.
314 Northeastern Naturalist Vol. 18, No. 3
Stehlik, L.L., and C.J. Meise. 2000. Diet of Winter Flounder in a New Jersey estuary:
Ontogenetic change and spatial variation. Estuaries 23(3):381–391.
Steimle, F.W., and R. Terranova. 1991. Trophodynamics of select demersal fishes in the
New York Bight. NOAA Technical Memorandum NMFS–F/NEC–84. National Marine
Fisheries Service, Woods Hole, MA. 11 pp.
Steimle, F.W., C.A. Zetlin, P.L. Berrien, D.L. Johnson, and S. Chang. 1999. Essential fish
habitat source document: Scup, Stenotomus chrysops, life history and habitat characteristics.
NOAA Technical Memorandum NMFS–NE–149. National Marine Fisheries
Service, Woods Hole, MA. 39 pp.
Steimle, F.W., R.A. Pikanowski, D.G. McMillan, C.A. Zetlin, and S.J. Wilk. 2000. Demersal
fish and American Lobster diets in the lower Hudson-Raritan estuary. NOAA
Technical Memorandum NMFS–NE161. National Marine Fisheries Service, Woods
Hole, MA. 106 pp.
US Fish and Wildlife Service (USFWS). 1997. Northeast coastal areas study significant
coastal habitats site 4 (NY). In Significant Habitats and Habitat Complexes of the
New York Bight Watershed. Available online at http://library.fws.gov/Pubs5/necas/
web_link/4_port%20jefferson.htm. Accessed 12 April 2010.
Wetherbee, B.M., and E. Cortés. 2004. Food consumption and feeding habits. Pp. 223–
244, In J.C. Carrier, J.A. Musick, and M.R. Heithaus (Eds.). Biology of Sharks and
their Relatives. CRC Press, Boca Raton, FL. 608 pp.
Wilcox, J.R., and H.P. Jeffries. 1973. Growth of the sand shrimp, Crangon septemspinosa,
in Rhode Island. Chesapeake Science 14(3):201−205.
Wilson, R.E., K.C. Wong, and H.H. Carter. 1991. Aspects of circulation and exchange in
Great South Bay. Pp. 9–22, In J.R. Schubel, T.M. Bell, and H.H. Carter (Eds.). The
Great South Bay. State University of New York Press, Stony Brook, NY. 107 pp.