Regular issues
Special Issues

Northeastern Naturalist
    NENA Home
    Aim and Scope
    Board of Editors
    Editorial Workflow
    Publication Charges

Co-published Journals
    Southeastern Naturalist
    Caribbean Naturalist

EH Natural History Home

Diet Composition and Feeding Habits of Common Fishes in Long Island Bays, New York
Skyler R. Sagarese, Robert M. Cerrato, and Michael G. Frisk

Northeastern Naturalist, Volume 18, Issue 3 (2011): 292–314

Full-text pdf (Accessible only to subscribers.To subscribe click here.)


Site by Bennett Web & Design Co.
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. Introduction 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 - 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 Study sites 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 Diet composition 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 (s2 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. Regional comparisons 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 Results 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). Diet composition 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 NENAonline/suppl-files/n18-3-972-Sagarese-s1, and, for BioOne subscribers, at 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://, and, for BioOne subscribers, at 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 s1, and, for BioOne subscribers, at 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. TL range 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 Teleosts 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 Crustaceans 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 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 Mollusks 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 Teleosts 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 Crustaceans 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 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 Mollusks 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. Regional comparisons 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). Discussion 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. Multivariate statistics 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. Acknowledgments 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. Literature Cited 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 Science 62:1076–1094. 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 234:191–204. 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 334:185–195. 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 A):41–54. 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 Series 217:157–165. Latour, R.J., M.J. Brush, and C.F. Bonzek. 2003. Toward ecosystem-based fisheries management: Strategies for multispecies modeling and associated data requirements. Fisheries 28:10–22. 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 106:47–57. Link, J.S. 2002. Ecological considerations in fisheries management: When does it matter? Fisheries 27(4):10–17. 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 57:697−706. 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 Bulletin 152:1–105. 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 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 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.