Technical Report FOOD HABITS OF YOUNG-OF-THE-YEAR ESTUARINE FISHES IN THE MIDDLE ATLANTIC BIGHT: A SYNTHESIS. Frank T. Mancini III and Kenneth W.

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1 Technical Report FOOD HABITS OF YOUNG-OF-THE-YEAR ESTUARINE FISHES IN THE MIDDLE ATLANTIC BIGHT: A SYNTHESIS Frank T. Mancini III and Kenneth W. Able August 2005 Institute of Marine and Coastal Sciences Rutgers, The State University of New Jersey 71 Dudley Road New Brunswick, NJ This is IMCS Contribution #

2 FOOD HABITS OF YOUNG-OF-THE-YEAR ESTUARINE FISHES IN THE MIDDLE ATLANTIC BIGHT: A SYNTHESIS Frank T. Mancini III and Kenneth W. Able August 2005 Marine Field Station Institute of Marine and Coastal Sciences Rutgers, The State University of New Jersey Tuckerton, NJ This is IMCS Contribution #

3 TABLE OF CONTENTS Abstract... 1 Introduction... 2 Methods... 2 Source of Literature... 2 Results... 3 Characterization of available data... 3 Major categories of prey consumed... 4 Species-specific variation in food habits... 4 Family Carcharhinidae... 4 Mustelus canis... 4 Family Anguillidae... 5 Anguilla rostrata... 5 Family Congridae... 5 Conger oceanicus... 5 Family Clupeidae... 6 Alosa aestivalis... 6 Alosa mediocris... 6 Alosa pseudoharengus... 6 Alosa sapidissima... 6 Brevoortia tyrannus... 7 Clupea harengus... 7 Family Engraulidae... 8 Anchoa hepsetus... 8 Anchoa mitchilli... 8 Family Osmeridae... 9 Osmerus mordax... 9 Family Synodntidae... 9 Synodus foetens... 9 Family Gadidae... 9 Microgadus tomcod... 9 Pollachius virens... 9 Family Phycidae Urophycis chuss Urophycis regia Urophycis tenuis Family Ophidiidae Ophidion marginatum Family Batrachoididae Opsanus tau Family Belondae Strongylura marina Family Cyprinodontidae Cyprinodon variegatus i

4 Family Fundulidae Fundulus heteroclitus Fundulus luciae Fundulus majalis Lucania parva Family Poeciliidae Gambusia holbrooki Family Atherinidae Membras martinica Menidia beryllina Menidia menidia Family Gasterosteidae Apeltes quadracus Gasterosteus aculeatus Family Syngnathidae Hippocampus erectus Syngnathus fuscus Family Triglidae Prinonotus carolinus Prinonotus evolans Family Cottidae Myoxocephalus aenaeus Family Percichthyidae Morone americana Morone saxatilis Family Serranidae Centropristis striata Family Pomatomidae Pomatomus saltatrix Family Carangidae Caranx hippos Family Lutjanidae Lutjanus griseus Family Sparidae Stenotomus chrysops Family Sciaenidae Bairdiella chrysoura Cynoscion regalis Leiostomus xanthurus Menticirrhus saxatilis Micropogonias undulatus Pogonias cromis Family Chaetodontidae Chaetodon ocellatus ii

5 Family Mugilidae Mugil cephalus Mugil curema Family Sphyraenidae Sphyraena borealis Family Labridae Tautoga onitis Tautogolabrus adspersus Family Pholidae Pholis gunnellus Family Uranoscopidae Astroscopus guttatus Family Blenniidae Hypsoblennius hentz Family Ammodytidae Ammodytes americanus Family Gobiidae Gobionellus boleosoma Gobiosoma bosc Gobiosoma ginsburgi Family Stromateidae Peprilus triacanthus Family Scophthalmidae Scophthalmus aquosus Family Paralichthyidae Etropus microstomus Paralichthys dentatus Family Pleuronectidae Pseudopleuronectes americanus Family Soleidae Trinectes maculates Family Tetraodontidae Sphoeroides maculates Discussion Limitations of this study Conclusions Acknowledgments Literature Cited List of Tables List of Figures Table 1. Identification of prey ranking by young-of-the-year estuarine fish based on composite of studies examined Table 2a. Identification of prey ranking by young-of-the-year estuarine fish based on each study examined iii

6 Table 2b. Details of specific studies which identified prey by young-of-the year estuarine fish Table 3. Details of specific species which identified fish as prey by young-ofthe-year estuarine fishes Table 3b. Identification of fish prey ranking by young-of-the-year estuarine fish based on each study examined Table 4. Size of young-of-the-year fish analyzed for stomach contents in the Middle Atlantic Bight based on the available literature Figure 1. Frequency distribution of number of food habit studies by fish species Figure 2. Frequency distribution of number of fish stomach samples by fish species Figure 3. Species specific size distribution of young-of-the-year fishes examined from the studies included in this analysis iv

7 ABSTRACT Estuaries are often presumed to be important habitats for juvenile fishes, in part, because they provide abundant food. In an effort to begin to evaluate this hypothesis we synthesized the available information from over 400 published papers, journal articles, theses, and dissertations on the food habits of the young-of-the-year (YOY) of 70 fish species, from 42 families, from the Middle Atlantic Bight (Cape Cod to Cape Hatteras) estuaries. Of the total, 91 papers contained food habit information for 47 of the 70 species. All species examined were YOY <300 mm in length with the exception of Mustelus canis (smooth dogfish), which ranged from mm TL. A sample size of over 54,000 stomachs was included in this synthesis. In the process of attempting to fill in the gaps of our understanding of YOY fish data, we are examining several aspects of food habits: availability or lack of food habit studies for YOY, prey categories consumed, ontogenetic and geographic variation in similarity in diet between closely related species, and the role of piscivory. The 47 species for which there are data consumed a variety of prey items, spanning 47 categories, suggesting opportunistic feeding for YOY fishes in general. Diet varied from species to species. While some species showed a narrow diet, others consumed a broad variety of prey. Seven categories of invertebrates were very important prey items across species, including copepods, amphipods, mysids, decapod shrimp, and crabs, polychaetes and insects/arachrids. Fish were also an important prey item for several species; the most important fish consumed were Anchoa mitchilli, Menidia menidia and Fundulus heteroclitus. Ontogenetic changes in diet were evident in several species as well as diet variation between estuaries, within estuaries between salinity zones, as well as seasonally and inter-annually. 1

8 INTRODUCTION One of the frequently cited assumptions about the functional significance of estuaries to young-of-the-year (YOY) fishes is that estuarine habitats provide abundant food sources, (Boesch and Turner, 1984; Blaber, 1981;Yáñez-Arancibia et al., 1980; Blaber and Blaber, 1980; Livingston, 2003) which contribute to their presumed nursery value (Beck et al. 2001). By reviewing and synthesizing the available literature, we hope to answer and pose several questions: What species have been studied for food habits during their first year in Middle Atlantic Bight estuaries? What species are lacking YOY food habit studies within this study area? What are the general patterns of YOY food habits? Are diets similar between closely related species? What role does piscivory play in diet during the first year? Unfortunately, these questions have been seldom addressed. In an earlier work, we had attempted to summarize much of the information about the first year in the life of estuarine fishes in the Middle Atlantic Bight (Able and Fahay 1998). In this extensive treatment, we neglected to include information on food habits (Moser, 1999). In an attempt to rectify this oversight, we have reviewed the available literature for YOY fishes that occur in estuaries in the Middle Atlantic Bight. Further, we attempted to discern species-specific patterns in the type of prey taxa consumed with some emphasis on fish prey. Sources of Literature METHODS Over 400 sources were utilized for this synthesis of literature on food habits of young-ofthe year estuarine fish in Middle Atlantic Bight (i.e. from Cape Cod, Massachusetts to Cape Hatteras, North Carolina) estuaries. Several criteria were evaluated before a publication was included in the synthesis. First, species of interest included dominant species in estuaries in the study area, i.e., the 70 species treated in Able and Fahay (1998). Second, we required that each study examine food habits of fish within the size range typical of young-of-the-year (YOY) for that species within the Middle Atlantic Bight based on the size limits in Able and Fahay (1998). Third, it was necessary that each study had a minimum study size (i.e. > 18 individuals/species). A published study or unpublished thesis or dissertation that met all of the necessary criteria was then further examined for detailed information on predator species and size and type of prey items consumed. The examination also included some description of the study area and habitat including location, salinity, method of ranking prey, sample size and literature source. This information was then tabulated in three ways: 1) a composite summary, consisting of a single line of information summarizing the food habits for each species (Table 1); 2) another summary that provided more detailed information from each individual study on the food habits of each predator species by study (Table 2); and 3) a separate summary that focused on those predator species that had fed on fish prey (Table 3). In addition, another 350 papers on food habits of the target species are listed, but not included in the synthesis because they were either outside the study area, not clearly YOY, or had too few specimens. 2

9 One difficulty encountered was that there was no standardization of prey categories or taxonomic level between authors or studies; therefore, choosing prey categories was sometimes difficult. In general, if a prey item was mentioned in any of the studies we examined, it was added to our list of prey items. It was not feasible to present every prey item at the species level; therefore, so many of the closely related organisms were combined into broader taxonomic levels. Prey categories were added and revised as necessary while reviewing the literature until we arrived at a final listing of potential prey taxa (see tables). It was difficult to standardize the importance of different prey taxa because methods of data analysis and presentation of results were not standard across studies. As a compromise, the information we collected from each individual study was arranged into three categories: Important Prey, Minor/Rare Prey, and Absent. Where possible, the category assigned to each prey item was based on the original author s interpretation of the results of each study. If the study author(s) suggested a prey item was important, it was classified as Important Prey. All other prey items noted by the study were considered Minor/Rare Prey. Prey items not mentioned by the study were considered Absent. The evaluation of importance was restricted to three levels largely in an effort to simplify the different methods of analysis and presentation of results used by different authors. In studies were the author(s) did not suggest Important Prey items, we provided our own interpretation, such that a prey item was considered important if it comprised greater than 20% of the diet. Characterization of Available Data RESULTS We have reviewed over 400 published papers, including theses and dissertations that were published on the food habits of YOY fishes of 70 target species from 42 families in the Middle Atlantic Bight between Of the total number of papers examined, 91 papers contained food habit information for 47 (67% of the 70) species. Of the 23 species for which there was no food habit information, all had food habit data available from outside the study area. Based on those studies with data, we have examined that data for over 54,000. The number of studies varied between species with several species (Pseudopleuronectes americanus, Leiostomus xanthurus, Cynoscion regalis, Pomatomus saltatrix) having over 10 published papers on food habits (Fig. 1). The individual sample sizes varied between each study; some were represented by >3000 stomachs while most were represented by <400 stomachs (Fig. 2). Most fishes studied were <300 mm in length with the exception of Mustelus canis, which ranged from mm in length (Fig. 3). 3

10 Major Categories of Prey Consumed Based on the available literature, prey were divided into 47 categories (Table 1). Seven categories of invertebrate prey were dominant (Important Prey of >20% of species examined) across many species including copepods, amphipods, mysids, decapod shrimps, polychaetes, crabs and insects/arachnids. As an indication, copepods were found to be Important Prey for 33 (70% of those for which there is data available) species, amphipods for 27 (57%) species, mysids for 26 (55%) species, decapod shrimps for 21 (45%) species, polychaetes for 17 (36%) species, crabs for 13 (28%) species and insects/arachnids for 11 (23%) species. In addition, these same prey categories were commonly consumed by all species studied as indicated by their frequency of occurrence, which was 81%, 91%, 74%, 72%, 79%, 68% and 51%, respectively. Other prey categories that were also frequently encountered (consumed by >30% of species examined) include isopods (74%) species, bivalves (62%), gastropods (55%), ostracods (51%), invertebrate eggs (47%), nematodes (38%), cnidarians (34%), algae (32%) and cladocerans (32%). Detritus and sediment were also frequently encountered (62% each). Fish were also Important Prey for some species (Table 1). Fish were important prey for 16 (34%) species, and they occurred in the diet of 34 (72%) species. Fish eggs also were consumed frequently (38% of species studied). In a further attempt to understand the role of fish as prey we tabulated the occurrence of individual fish prey. Of the 34 species that consumed fish prey, the data for 24 species included sufficient information to characterize the fish species consumed (Table 3). Some of the dominant (important to >15% of species examined) fish prey were Fundulus heteroclitus (Important Prey for 21% of species), Menidia menidia (17%), and Anchoa mitchilli (17%). These same species were found in 29%, 29% and 42% of the species examined, respectively. Other fish species that were frequently encountered (consumed by >13% of species examined) in stomach contents were Menidia spp., Anchoa spp., Family Gobiidae, Gobiosoma bosc, Gobiosoma spp., Family Clupeidae, Micropogonias undulatus, Ammodytes americanus, Family Atherinopsidae, Alosa spp.). Species-Specific Variation in Food Habits Mustelus canis, smooth dogfish Family Carcharhinidae One study examined the food habits of YOY ( mm TL, n= 85) smooth dogfish, Mustelus canis, in the Barnegat Bay - Little Egg Harbor and the Mullica River Great Bay estuaries, New Jersey (Rountree and Able 1996) (Tables 1, 2, 3). Important Prey of YOY included a variety of invertebrates (polychaetes, decapod shrimps, crabs). YOY also consumed bivalves and small fish (Menidia menidia, Fundulus heteroclitus), although less frequently than invertebrates. Several other papers have been published on food habits of smooth dogfish, but they are not within the geographical range/age scope of this study (Breder 1921; Bigelow and 4

11 Schroeder 1953; Kimmel 1973; Gelsleichter et al. 1999; Bowman et al. 2000; Branstetter 2002). Anguilla rostrata, American eel Family Anguillidae Several studies (n=3) have examined the food habits of YOY ( mm TL, n=187) American eel, Anguilla rostrata, in Middle Atlantic Bight estuaries from New Jersey streams to the James River, Virginia (Tables 1, 2, 3). Important Prey of YOY were invertebrates (insects/arachnids, Limulus spp. eggs and larvae). Fish (Menidia menidia) were a minor dietary component. The diet of YOY shifted with ontogeny at approximately mm TL, such that insects were Important Prey of small eels and fish and crayfish were Important Prey of larger eels. Ogden (1970) noted a shift in diet with ontogeny, such that eels <400 mm TL contained primarily bottom dwelling insects, while larger eels contained larger prey items, almost exclusively crayfish and fish. Lookabaugh and Angermeier (1992) suggested a similar transition but at approximately 250 mm TL. Variation in diet has also been noted within the James River drainage, Virginia; with stomachs of YOY ( mm TL, n=20) from the upper Piedmont region containing almost exclusively aquatic invertebrates, while YOY ( mm TL, n=19) from the lower Piedmont contained some terrestrial invertebrates and crayfish in addition to aquatic invertebrates and YOY ( mm TL, n=17) from the coastal plain contained some crayfish in addition to aquatic invertebrates (Lookabaugh and Angermeier 1992). Several other papers have been published on food habits of American eel, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Brinley and Bowen 1935; Vladykov 1955; Godfrey 1957; Wenner and Musick 1975; Facey and LaBar 1981; Sorenson et al. 1986; Denoncourt and Stauffer 1993; Smith and Tighe 2002a), or were unavailable (Compton 1968). Conger oceanicus, conger eel Family Congridae There have been no studies on the food habits of YOY (<350 mm) Conger eel, Conger oceanicus, in Middle Atlantic Bight estuaries. However, several other papers have been published on food habits, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Maurer and Bowman 1975; Cau and Manconi 1984; Levy et al. 1988; Bowman et al. 2000; Smith and Tighe 2002b). 5

12 Family Clupeidae Alosa aestivalis, blueback herring Several studies (n=4) have examined the food habits of YOY (5-106 mm, n=1610) blueback herring, Alosa aestivalis, in Middle Atlantic Bight estuaries from the Connecticut River to James River, Virginia (Tables 1, 2). Important Prey of YOY included a variety of invertebrates (copepods, cladocerans, insects/arachnids, rotifers). These studies do not show a shift in diet with ontogeny or over the geographical range. Several other papers have been published on food habits of blueback herring, but they are not within the geographical range/age scope of this study (Hildebrand 1963; Richards 1963; Holland and Yelverton 1973; Maurer and Bowman 1975; Creed 1985; Stone 1986; Stone and Daborn 1987; Scott and Scott 1988; Bowman et al. 2000; Munroe 2002a), or were unavailable (Davis and Cheek 1966; Frankensteen 1976; Domermuth 1976). Alosa mediocris, hickory shad There have been no studies of the food habits of YOY (<150 mm) hickory shad, Alosa mediocris, in Middle Atlantic Bight estuaries. However, several other papers have been published on food habits, but they are not within the geographical range/age scope of this study (Hildebrand 1963; Bowman et al. 2000; Munroe 2002a), or were unavailable (Frankensteen 1976). Alosa pseudoharengus, alewife One study examined the food habits of YOY (mean 64.8 mm TL, n=48) alewife, Alosa pseudoharengus, in the Hudson River estuary (Grabe 1996) (Tables 1, 2). YOY fed primarily on chironomid insects and amphipods. Several other papers have been published on food habits of alewife, but they are not within the geographical range/age scope of this study (Hildebrand 1963; Richards 1963; Edsall 1964; Morsell and Norden 1968; Norden 1968; Lackey 1969; Hutchinson 1971; Holland and Yelverton 1973; Vigerstad and Cobb 1978; Nigro and Ney 1982; Johnson 1983; Gregory et al. 1983; Stone and Jessop 1994; Mills et al. 1995; Bowman et al. 2000; Munroe 2002a), or were unavailable (Weaver 1975; Frankensteen 1976). Alosa sapidissima, American shad Several studies (n=4) have examined the food habits of YOY (10-95 mm FL, n=2861) American shad, Alosa sapidissima, in Middle Atlantic Bight estuaries from Connecticut River to Neuse River, North Carolina (Tables 1, 2, 3). Important Prey of YOY included a variety of invertebrates (insects/arachnids, copepods, ostracods, nematodes) and fish (cyprinids, Anchoa spp.). These studies do not show a shift in diet with ontogeny or over the geographical range. 6

13 Several other papers have been published on food habits of American shad, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Atkinson 1951; Bigelow and Schroeder 1953; Hildebrand 1963; Cheek 1968; Levesque and Reed 1972; Williams and Bruger 1972; Holland and Yelverton 1973; Kimmel 1973; Marcy 1976b; Domermuth and Reed 1980; Bowman et al. 2000; Munroe 2002a), or were unavailable (Domermuth 1976; Frankensteen 1976). Brevoortia tyrannus, Atlantic menhaden Several studies (n=6) have examined the food habits of YOY ( mm, n>719) Atlantic menhaden, Brevoortia tyrannus, in Middle Atlantic Bight estuaries from Narragansett, Rhode Island to Newport River, North Carolina (Tables 1, 2, 3). Important Prey of YOY included detritus and a variety of small invertebrates (copepods, zooplankton, mysids, cirripedes) and phytoplankton (diatoms, flagellates, algae). Fish (Anchoa mitchilli) were a minor dietary component. The diet of YOY shifted with ontogeny from a diet dominated by zooplankton at smaller sizes to one dominated by phytoplankton at larger sizes. Small YOY (19-34 mm FL, n=243) consumed almost exclusively zooplankton (copepods), whereas gut contents of progressively larger YOY (28-94 mm FL, n=117) shifted to a mixture of detritus, diatoms and flagellates (June and Carlson 1971). However, there was no change in diet with size of mm YOY (n=300) (Mulkana 1964). Geographical variation in diet has also been noted within estuaries with some YOY (40-60 mm, n=30-40) relying less on zooplankton and more on suspended organic matter from bay through river to marsh in Narragansett Bay, Rhode Island (Jeffries 1975). Several other papers have been published on food habits of Atlantic menhaden, but they are not within the geographical range/age scope of this study (Peck 1894; Hildebrand and Schroeder 1928; Bigelow and Schroeder 1953; Darnell 1958; Hildebrand 1963; Kimmel 1973; Peters and Kjelson 1975; Durbin and Durbin 1975; Edgar and Hoff 1976; Bowman et al. 2000; Munroe 2002a). Clupea harengus, Atlantic herring One study examined the food habits of YOY ( mm, n=18) Atlantic herring, Clupea harengus, in Long Island Sound (Richards 1963) (Tables 1, 2). Copepods were the bulk of the diet, but mysids, diatoms and dinoflagellates were also consumed. Several other papers have been published on food habits of Atlantic herring, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Battle et al. 1936; Legare and MacClellan 1960; Hildebrand 1963; Sherman and Honey 1968; Sherman and Perkins 1971; Maurer 1976; Cohen and Lough 1983; James 1988; Blaxter 1990; Bowman et al. 2000; Munroe 2002a). 7

14 Family Engraulidae Anchoa hepsetus, striped anchovy There have been no studies of the food habits of YOY (<100 mm) striped anchovy, Anchoa hepsetus, in Middle Atlantic Bight estuaries. However, several other papers have been published on food habits, but they are not within the geographical range/age scope of this study (Hildebrand and Cable 1930; Stevenson 1958; Springer and Woodburn 1960; Hildebrand 1963; Carr and Adams 1973; Adams 1976; Modde and Ross 1983; DeLancey 1989; Allen et al. 1995; Bowman et al. 2000; Munroe 2002a). Anchoa mitchilli, bay anchovy Several studies (n=6) have examined the food habits of YOY ( mm, n=6179) bay anchovy, Anchoa mitchilli, in Middle Atlantic Bight estuaries from Delaware Bay to Newport River, North Carolina (Tables 1, 2, 3). Important Prey of YOY included a variety of invertebrates (copepods, mysids, crabs, ostracods). Fish (Anchoa mitchilli, Micropogonias undulatus) were a minor dietary component with instances of cannibalism (Nemerson 2001). The diet of YOY shifted with ontogeny from a diet of various small invertebrates (copepods, crabs, ostracods) to one dominated by mysids supplemented by other invertebrates and fish with increasing size. The diet of YOY ( mm, n=1485) larvae from Chesapeake Bay also shifted to include greater amounts of copepods with increasing size, such that YOY mm fed on copepod eggs and nauplii, rotifers, tintinnids and diatoms, while YOY mm fed primarily on copepodites and adult copepods (78% F) (Auth 2003). In Delaware Bay, the diet of YOY (10-60 mm, n=3227) shifted at approximately 50 mm from zooplankton to mysids, crustaceans and fish with growth (Nemerson 2001). The diet shifted earlier at approximately 20 mm in York River, Virginia, such that the dominant prey of YOY mm was crabs, ostracods, and copepods, while the dominant prey of YOY mm was mysids supplemented by various invertebrates and detritus (Smith et al. 1984). The diet suggested by these studies was consistent over the geographic range. Relative importance of individual prey items also varied intra-annually in Delaware Bay (Nemerson 2001) and inter-annually in Chesapeake Bay (Auth 2003). Several other papers have been published on food habits of bay anchovy, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Darnell 1958; Springer and Woodburn 1960; Hildebrand 1963; Richards 1963; Odum 1971; Odum and Heald 1972; Carr and Adams 1973; Kimmel 1973; Adams 1976; Sheridan 1978; Kinch 1979; Din and Gunter 1986; Vazquez Rojas 1989; DeLancey 1989; Johnson et al. 1990; Allen et al. 1995; Munroe 2002a), or were unavailable (Van Engle and Joseph 1968; Klebasko 1991). 8

15 Family Osmeridae Osmerus mordax, rainbow smelt There have been no studies of the food habits of YOY (<75 mm) rainbow smelt, Osmerus mordax, in Middle Atlantic Bight estuaries. However, several other papers have been published on food habits, but they are not within the geographical range/age scope of this study (Gordon 1961; Bigelow and Schroeder 1953; Richards 1963; Lackey 1969; Siefert 1972; Scott and Crossman 1973; Foltz 1974; O'Gorman 1974; McCullough and Stanley 1981; Scott and Scott 1988; Urban and Brandt 1993; Mills et al. 1995; Klein-MacPhee 2002b), or were unavailable (Price 1963). Synodus foetens, inshore lizardfish Family Synodntidae There have been no studies of the food habits of YOY (<250 mm) inshore lizardfish, Synodus foetens, in Middle Atlantic Bight estuaries. However, several other papers have been published on food habits, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Beebe and Tee-Van 1933; Reid 1954, 1955; Springer and Woodburn 1960; Randall 1967; Carr and Adams 1973; Phelps 1997; Bowman et al. 2000). Microgadus tomcod, Atlantic tomcod Family Gadidae Several studies (n=2) have examined the food habits of YOY ( mm, n>557) Atlantic tomcod, Microgadus tomcod, in the Hudson River estuary of the Middle Atlantic Bight (Tables 1, 2). Important Prey of YOY included a variety of small invertebrates (amphipods, copepods, mysids, isopods). The diet of YOY ( mm, n= 577) shifted with ontogeny in Haverstraw Bay, New York, such that the importance of copepods decreased and amphipod importance increased as length increased, with amphipods dominant by 90 mm (Grabe 1978). Relative importance of individual prey items also varied intra-annually, copepods were dominant during May-June, whereas amphipods, mysids and isopods were dominant from July-December (Grabe 1978). Several other papers have been published on food habits of Atlantic tomcod, but they are not within the geographical range/age scope of this study (Alexander 1971; Lambert and Fitzgerald 1979; Grabe 1980; Klein-MacPhee 2002c; Able and Duffy-Anderson 2005). Pollachius virens, pollock One study examined the food habits of YOY ( mm, n=22) pollock, Pollachius virens, in Long Island Sound (Richards 1963) (Tables 1, 2). YOY consumed principally 9

16 copepods and mysids. Mysids were most frequent in fish mm, while larger fish (55-66 mm) also consumed amphipods. Several other papers have been published on food habits of pollock, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Marak 1960; Langton and Bowman 1980; Bowman 1981; Bowman and Michaels 1984; Bowman et al. 1987; Ojeda and Dearborn 1991; Bromley et al. 1997; Bowman et al. 2000; Klein-MacPhee 2002c). Urophycis chuss, red hake Family Phycidae One study examined the food habits of YOY ( mm, n=48) red hake, Urophycis chuss, in Long Island Sound (Richards 1963) (Tables 1, 2). Decapod shrimp were the basis of the diet of YOY supplemented by small proportions of other invertebrates. Several other papers have been published on food habits of red hake, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Vinogradov 1972; Langton and Bowman 1980; Hacunda 1981; Bowman 1981; Luczkovich 1982; Sedberry 1983; Luczkovich and Olla 1983; Garman 1983; Bowman and Michaels 1984; Bowman et al. 1987; Garrison and Link 2000; Bowman et al. 2000; Klein-MacPhee 2002c). Urophycis regia, spotted hake Two studies examined the food habits of YOY ( mm, n=78) spotted hake, Urophycis regia, in Long Island Sound and Magothy Bay, Virginia (Tables 1, 2, 3). Important Prey of YOY included a variety of invertebrates (decapod shrimps, amphipods, mysids). Fish were a minor dietary component, mainly Ammodytes americanus in Long Island Sound (Richards 1963) and Anchoa mitchilli in Magothy Bay (Kimmel 1973). In Magothy Bay, smaller YOY (34-80 mm, n=26) consumed large percentages of smaller food items (amphipods, mysids, copepods, ostracods), while larger YOY ( mm, n=19) consumed larger items including decapod shrimp (Crangon septemspinosa) and fish (Anchoa mitchilli) (Kimmel 1973). Several other papers have been published on food habits of spotted hake, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Bigelow and Schroeder 1953; Barans 1969; Sikora et al. 1972; Langton and Bowman 1980; Sedberry 1983; Bowman and Michaels 1984; Burr and Schwartz 1986; Bowman et al. 1987; Garrison and Link 2000; Bowman et al. 2000; Klein-MacPhee 2002c). 10

17 Urophycis tenuis, white hake There have been no studies of the food habits of YOY (<250 mm) white hake, Urophycis tenuis, in Middle Atlantic Bight estuaries However, several other papers have been published on food habits, but they are not within the geographical range/age scope of this study (Bowman 1981; Coates et al. 1982; Bowman et al. 1987; Garrison and Link 2000; Bowman et al. 2000; Klein-MacPhee 2002c). Ophidion marginatum, striped cusk-eel Family Ophidiidae There have been no studies of the food habits of YOY (<200 mm) striped cusk-eel, Ophidion marginatum, in Middle Atlantic Bight estuaries. However, several other papers have been published on food habits, but they are not within the geographical range/age scope of this study (Kimmel 1973; Schwartz 1997b; Bowman et al. 2000; Collette and Klein-MacPhee 2002b). Opsanus tau, oyster toadfish Family Batrachoididae One study examined the food habits of YOY ( mm TL, n=56) oyster toadfish, Opsanus tau, in Chesapeake Bay (Chrobot, 1951) (Tables 1, 2, 3). The principal food items were arthropods, crabs (Eurypanopeus depressus, Callinectes sapidus), decapod shrimp (Palaemonetes vulgaris) and amphipods. Fishes (Apeltes quadracus, Menidia menidia, M.beryllina, Fundulus majalis, Lucania parva, Gobiosoma bosc) were a minor part of the diet. YOY also exhibited cannibalistic behavior. The diet of YOY shifted with ontogeny at approximately 100 mm TL, from a diet of invertebrates (crabs, arthropods, amphipods, decapod shrimp) to a diet of crabs and fish. YOY (<100 mm TL) consumed mud crabs, grass shrimp, amphipods and miscellaneous arthropods, whereas larger fish consumed primarily crabs and fish. Relative importance of individual prey items also varied intra-annually. In May YOY consumed mostly fish with a small percentage of crabs. From July to August there was an abundance of both fish and crabs, as well as numerous arthropods, amphipods, and shrimp. Crabs were the only abundant food item in September. Several other papers have been published on food habits of oyster toadfish, but they are not within the geographic range/age scope of this study (Gudger 1910; Hildebrand and Schroeder 1928; Bigelow and Schroeder 1953; Schwartz and Dutcher 1963; McDermott 1965; Kimmel 1973; Adams 1976; Wilson et al. 1982; Timmons 1995; Collette 2002a). 11

18 Strongylura marina, Atlantic needlefish Family Belonidae There have been no studies of the food habits of YOY (<400 mm) Atlantic needlefish, Strongylura marina, in Middle Atlantic Bight estuaries. However, several other papers have been published on food habits, but they are not within the geographical range/age scope of this study (McLane 1955; Carr and Adams 1973; Bowman et al. 2000; Collette 2002c). Cyprinodon variegatus, sheepshead minnow Family Cyprinodontidae Several studies (n=3) have examined the food habits of YOY (10-57 mm SL, n=1147) sheepshead minnow, Cyprinodon variegatus, in Middle Atlantic Bight estuaries from Hereford Inlet, New Jersey to Indian River Bay, Delaware (Tables 1, 2). Important Prey of YOY included phytoplankton (algae, diatoms, foraminiferans, flagellates) and a variety of invertebrates (nematodes, polychaetes, amphipods, copepods), as well as detritus and sediment. Fish were a minor dietary component. At Hereford Inlet, New Jersey the diet of YOY (10-35 mm SL, n=50) shifted with ontogeny at approximately 20 mm, from a diet of meiobenthic animals to a more herbivorous diet (Clymer 1978). YOY (<20 mm) consumed primarily invertebrates (copepods, polychaetes, nematodes), whereas larger YOY consumed primarily phytoplankton (algae, diatoms, foraminiferans) and detritus. Several other papers have been published on food habits of sheepshead minnow, but they are not within the geographical range/age scope of this study (Springer and Woodburn 1960; Harrington and Harrington 1961; Martin 1970; Odum 1971; Odum 1972; Kimmel 1973; Stickney and McGeachin 1975). Fundulus heteroclitus, mummichog Family Fundulidae Several studies (n=6) have examined the food habits of YOY ( mm SL, n=828) mummichog, Fundulus heteroclitus, in Middle Atlantic Bight estuaries from Hereford Inlet, New Jersey to Newport River, North Carolina (Tables 1, 2, 3). Important Prey of YOY included a variety of invertebrates (copepods, amphipods, polychaetes, tanaids, insects, oligochaetes) and phytoplankton (algae, diatoms). Fish (Fundulus spp.) and eggs were minor dietary components. The diet of YOY shifted with ontogeny at approximately 30 mm, from largely carnivorous at smallest sizes (amphipods, copepods, tanaids) to more omnivorous (detritus, algae, crabs, insects, polychaetes) at larger sizes (Schmelz 1964; Kneib and Stiven 1978; Morgan 1990; Smith et al. 2000). However, in 12

19 Hereford Inlet, New Jersey YOY remained carnivorous at larger sizes (16-40 mm, n= 50) (Clymer 1978). Relative importance of individual prey items also varied intra-annually in North Carolina salt marshes, such that algae, insects, crabs and detritus were consumed more frequently in summer and fall, while small crustaceans and polychaetes were frequently consumed during the entire year (Kneib and Stiven 1978). Several other papers have been published on food habits of mummichog, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Bigelow and Schroeder 1953; de Sylva et al. 1962; Kimmel 1973; Fritz 1974; Stickney and McGeachin 1975; Baker-Dittus 1978; Kneib 1986; Allen et al. 1994; Fell et al. 1998; James-Pirri et al. 2001; Able 2002). Fundulus luciae, spotfin killifish Two studies have examined the food habits of YOY (11-47 mm, n=370) spotfin killifish, Fundulus luciae, in Middle Atlantic Bight estuaries at York River, Virginia and Bogue Sound, North Carolina (Tables 1, 2). Important Prey of YOY included detritus, phytoplankton (diatoms) and a variety of invertebrates (ostracods, insects, copepods, tanaids). At Fox Creek Marsh, Virginia, fish eggs were a minor dietary component (Byrne 1978). Relative importance of individual prey items varied intra-annually at this same site (Byrne 1978) as well as in Tar Landing Bay, North Carolina, where summer and fall diet was dominated by organic matter (detritus, diatoms, sand), while winter and spring diet was dominated by small crustaceans (copepods, tanaids, amphipods) and insects (Kneib 1978). One other paper has been published on food habits of the spotfin killifish (Hildebrand and Schroeder 1928). Fundulus majalis, striped killifish Two studies have examined the food habits of YOY (26-70 mm SL, n=62) striped killifish, Fundulus majalis, in Middle Atlantic Bight estuaries at Hereford Inlet, New Jersey and Delaware River estuary (Tables 1, 2). Important Prey of YOY included a variety of invertebrates (copepods, nematodes, Limulus spp. eggs and larvae). Fish were a minor dietary component in the Delaware River estuary (de Sylva et al. 1962). Several other papers have been published on food habits of striped killifish, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Bigelow and Schroeder 1953; Kimmel 1973; Harvey 1998; Able 2002), or were unavailable (Werme, 1981; Petrillo 1987). Lucania parva, rainwater killifish There have been no studies of the food habits of YOY (<40 mm TL) rainwater killifish, Lucania parva, in Middle Atlantic Bight estuaries. However, several other papers have 13

20 been published on food habits, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Simpson and Gunter 1956; Harrington and Harrington 1961; Odum 1971; Kimmel 1973; Moyle 1976). Gambusia holbrooki, eastern mosquitofish Family Poeciliidae There have been no studies of the food habits of YOY (<40 mm TL) eastern mosquitofish, Gambusia holbrooki, in Middle Atlantic Bight estuaries. However, one other paper has been published on food habits, but it is not within the geographical range/age scope of this study (Garcia-Berthou 1999). Membras martinica, rough silverside Family Atherinidae There have been no studies of the food habits of YOY (<110 mm FL) rough silverside, Membras marinica, in Middle Atlantic Bight estuaries. However, several other papers have been published on food habits, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Reid 1954; Stickney and McGeachin 1975; Burton 1982; Allen et al. 1995). Menidia beryllina, inland silverside Several studies (n=3) have examined the food habits of YOY (15-67 mm TL, n=522) inland silverside, Menidia beryllina, in Middle Atlantic Bight estuaries from Great South Bay, New York to Barnegat Bay, New Jersey (Tables 1, 2). Important Prey of YOY included a variety of invertebrates (copepods, rotifers, insects, crabs), phytoplankton (algae), detritus, fish and fish eggs. These studies showed variation in diet between estuaries, such that YOY in Great South Bay, New York relied on decapod larvae, fish eggs and larvae (Grover 1982), while insects, rotifers and detritus were Important Prey items of YOY in New Jersey salt marshes (Coorey 1981; Coorey et al. 1985). Copepods were Important Prey items in both estuaries. Geographical variation in diet has also been noted between salt marshes within the same New Jersey estuary (Coorey 1981). At oligo-polyhaline Cattus Island, copepods (46.7% composition) were most important to diet of YOY (15-40 mm TL, n=218), supplemented by rotifers, detritus and algae (19.1, 20.8, 12.4%), while insects (60.9%) were most important to YOY (15-40 mm TL, n=6) at mesohaline Popular Point, supplemented by copepods and rotifers (16.8, 5.8%). However, detritus and algae were not consumed. Relative importance of individual prey items also varied intra-annually in Great South Bay, New York, such that copepods decreased in percent composition of diet from spring to summer, then were the only prey consumed in fall, while decapod larvae and fish eggs and larvae became more important in summer (Grover 1982). Similar seasonal trends were noted at Cattus Island and Cedar Run, New Jersey, where copepods dominated diet during the entire year and algae, and 14

21 rotifers were utilized during summer and fall months (Coorey 1981). These studies did not show an ontogenetic shift in diet during the first year. Several other papers have been published on food habits of inland silverside, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Darnell 1958; Carr and Adams 1973; Stickney and McGeachin 1975; Bengston 1984; Bowman et al. 2000; Chernoff 2002), or were unavailable (Reid 1954). Menidia menidia, Atlantic silverside Several studies (n=7) have examined the food habits of YOY ( mm, n=1439) Atlantic silverside, Menidia menidia, in Middle Atlantic Bight estuaries from Lower Pettaquamscutt River, Rhode Island to Newport River, North Carolina (Tables 1, 2 and 3). Important Prey of YOY included a variety of invertebrates (rotifers, polychaetes, insects, mysids, amphipods, decapod shrimps, crabs, copepods, cirripedes, Limulus spp., eggs), fish (Menidia menidia, Fundulus spp., eggs), phytoplankton (diatoms) and detritus. YOY exhibit cannibalistic behavior (Cadigan and Fell 1985). The diet of YOY shifted with ontogeny (between mm), from a diet of small invertebrates (copepods, cirripedes, insects), invertebrate eggs and phytoplankton (diatoms) to a diet of larger invertebrates (decapod shrimp, mysids, amphipods, isopods) and fish with increasing size (Mulkana 1964; Kimmel 1973; Cadigan and Fell 1985; Morgan 1990). In the Pataguanset estuary, Connecticut, the diet of YOY ( mm, n=147) shifted at approximately 80 mm, such that smaller fish most frequently ate copepods, barnacle nauplii, and insects, while larger individuals preferred shrimp, fish and amphipods (Cadigan and Fell 1985). Morgan (1990) found results similar to these in the Newport River estuary, North Carolina. In Magothy Bay, Virginia, smaller YOY (16-69 mm, n=19) consumed higher percentages of copepods, while larger fish ( mm, n=28) consumed higher percentages of mysids (Kimmel 1973). However, in Lower Pettaquamscutt River, Rhode Island, the diet shifted at approximately 30 mm. Important food components for small YOY (10-30 mm, n=69) included diatoms, rotifers, barnacle nauplii, and invertebrate eggs; Important Prey items of larger YOY (31-80 mm, n=81) included amphipods, isopods and insects, while copepods of various stages were Important Prey to all sizes examined (Mulkana 1964). Spatial variation in diet has also been noted within Pataguanset estuary, Connecticut, such that shrimp, copepods and eggs were dominant prey in the lower estuary, while plant material, fish and copepods were most abundant in the upper estuary (Cadigan and Fell 1985), and in Narragansett, Rhode Island, where YOY from lower Pettaquamscutt River consumed a mixture of plant and animal food, while YOY from lower Point Judith Pond were largely dependent upon phytoplankton (Mulkana 1964). Relative importance of individual prey items also varied intra-annually in Magothy Bay, Virginia, such that during winter (December-March) mysids and polychaetes were the principal prey, while amphipods and calanoid copepods were the main diet during spring and summer. Seasonal diet variation was also noted in the Pataguanset River estuary, Connecticut (Cadigan and Fell 1985) and Great South Bay, New York (Grover 1982). 15

22 Several other papers have been published on food habits of Atlantic silverside, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Bigelow and Schroeder 1953; Adams 1976; Gilmurray and Daborn 1981; Burton 1982; Bengston 1984; Scott and Scott 1988; Warkentine and Rachlin 1989; DeLancey 1989; Allen et al. 1995; Bowman et al. 2000; Chernoff 2002), or were unavailable (Bayliff 1950; Spraker and Austin 1997). Apeltes quadracus, fourspine stickleback Family Gasterosteidae There have been no studies of the food habits of YOY (<60 mm TL) fourspine stickleback, Apeltes quadracus in Middle Atlantic Bight estuaries. However, several other papers have been published on food habits, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Richards 1963; Delbeek and Williams 1987, 1988; Krueger 2002). Gasterosteus aculeatus, threespine stickleback There have been no studies of the food habits of YOY (<50 mm) threespine stickleback, Gasterosteus aculeatus, in Middle Atlantic Bight estuaries However, several other papers have been published on food habits, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Hynes 1950; Richards 1963; Walsh and FitzGerald 1984; Delbeek and Williams 1987; Foster et al. 1988; Delbeek and Williams 1988; Poulin and FitzGerald 1989; Bowman et al. 2000; Krueger 2002). Hippocampus erectus, lined seahorse Family Syngnathidae One study examined the food habits of YOY ( mm TL, n=136) lined seahorse, Hippocampus erectus, in lower York River, Virginia (Teixeira 1995) (Tables 1 and 2). Amphipods were the most Important Prey followed by copepods. The diet of YOY shifted with ontogeny at approximately 60 mm, such that YOY <60 mm TL fed mainly on amphipods and secondarily on copepods, whereas YOY >60 mm fed almost exclusively on amphipods. Several other papers have been published on food habits of lined seahorse, but they are not within the geographical range/age scope of this study (Bigelow and Schroeder 1953; Kimmel 1973; Howells and Howells 1989; Klein-MacPhee 2002e). Syngnathus fuscus, northern pipefish Several studies (n=4) have examined the food habits of YOY ( mm TL, n=5542) 16

23 northern pipefish, Syngnathus fuscus, in Middle Atlantic Bight estuaries from Long Island Sound, New York to lower York River, Virginia (Tables 1, 2,3). Important Prey of YOY included a variety of invertebrates (amphipods, copepods, isopods, mysids, cladocerans, gastropods). Fish (Syngnathus fuscus, Hippocampus erectus) were a minor dietary component. YOY exhibit cannibalistic behavior on a minor scale (Teixeira 1995). The diet of YOY shifted with ontogeny at approximately 100 mm in Chesapeake Bay, from a diet of predominantly copepods at smaller sizes, to amphipods and isopods when larger (Ryer and Orth 1987; Teixeira 1995). Relative importance of individual prey items also varied intra-annually in the lower Chesapeake Bay, such that gammarid and caprellid amphipods and calanoid copepods dominated the diet in spring and summer, isopods dominated diet during fall, while mysids and calanoid copepods dominated winter diet (Ryer and Orth 1987). Several other papers have been published on food habits of northern pipefish, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Bigelow and Schroeder 1953; Kimmel 1973; Adams 1976; Howard and Koehn 1985; Bowman et al. 2000; Klein-MacPhee 2002e). Prionotus carolinus, northern searobin Family Triglidae Several studies (n=4) have examined the food habits of YOY ( mm TL, n=295) northern searobin, Prionotus carolinus, in Middle Atlantic Bight estuaries from Woods Hole, Massachusetts to Magothy Bay, Virginia (Tables 1, 2 and 3). Important Prey of YOY included a variety of invertebrates (mysids, copepods, amphipods, decapod shrimps, polychaetes, cumaceans, isopods, crabs) and fish (Anchoa mitchilli). Sediment was also prominent in stomachs. In Long Island Sound, the diet of YOY ( mm SL, n=120) shifted with ontogeny such that small YOY consumed copepods, and as they grew through the fall, their diet changed to amphipods, isopods, cumaceans and small crabs (Richards et al. 1979). Similarly, in Magothy Bay, Virginia, smaller (22-54 mm, n=24) YOY fed mainly on Neomysis americana (82.2% volume) supplemented by amphipods (10.1%) and small Crangon septemspinosa (4.7%), while larger ( mm, n=24) YOY did not feed as heavily on N.americana (44.5%), but increased consumption of amphipods (16.3%), decapod shrimps (12.5%), isopods (8.7%), crabs (4.0%) and plant material. Several other papers have been published on food habits of northern searobin, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Bigelow and Schroeder 1953; Roberts 1978; Bowman et al. 2000; Klein-MacPhee and McBride 2002). Prionotus evolans, striped searobin Two studies examined the food habits of YOY ( mm SL, n=309) striped searobin, 17

24 Prionotus evolans, in Woods Hole, Massachusetts and New Haven and the Connecticut shore of Long Island Sound (Tables 1 and 2). Important Prey of YOY included a variety of invertebrates (decapod shrimps, copepods, gastropods, bivalves, mysid shrimps, eggs), fish and fish eggs. In Long Island Sound, diet of YOY ( mm SL, n=271) shifted with ontogeny, such that small YOY consumed copepods and larger sizes in the fall consumed more Neomysis americana, Crangon septemspinosa, a few mollusks and fish (Richards et al. 1979). Several other papers have been published on food habits of striped searobin, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Richards 1963; Kimmel 1973; Roberts-Goodwin 1981; Manderson et al. 1999; Bowman et al. 2000; Klein-MacPhee and McBride 2002). Myoxocephalus aenaeus, grubby Family Cottidae Two studies examined the food habits of YOY ( , n=196) grubby, Myoxocephalus aenaeus, in Cape Cod, Massachusetts and on the Connecticut shore of Long Island Sound (Tables 1, 2, 3). Important Prey of YOY included a variety of invertebrates (decapod shrimps, mysids, amphipods, isopods) and fish (Fundulus heteroclitus). In Nauset Marsh estuary, Massachusetts the diet of YOY ( mm SL, n=168) shifted with ontogeny at approximately 60 mm, from a diet of predominantly crustaceans (Crangon septemspinosa, isopods, amphipods) when small, to a diet of crustaceans (C. septemspinosa, Carcinus maenas) and fish (F. heteroclitus, Apeltes quadracus) when larger (Lazzari et al. 1989). Several other papers have been published on food habits of grubby, but they are not within the geographical range/age scope of this study (Laroche 1982; Klein-MacPhee 2002f). Morone americana, white perch Family Percichthyidae Several studies (n=6) have examined the food habits of YOY ( mm, n=2971) white perch, Morone americana, in Middle Atlantic Bight estuaries, from the Connecticut River to the York River, Virginia (Tables 1, 2, 3). Important Prey items of YOY included a variety of invertebrates (zooplankton, nemertines, polychaetes, oligochaetes, insects/arachnids, mysids, cumaceans, isopods, amphipods, decapod shrimps, crabs, ostracods, copepods, cladocerans, invertebrate eggs, detritus) and fish (Anguilla rostrata, Notropis hudsonius, Morone americana) including incidences of cannibalism (Marcy 1976a). The diet of YOY shifted with ontogeny at approximately mm, becoming more diverse with increased size (Miller 1963; Marcy 1976a; Smith et al. 1984; Bath and O Connor 1985). Marcy (1976a) found YOY (<100 mm TL, 18

25 n=759) fed less on plankton (copepods, cladocerans, amphipods) and more on benthic organisms (oligochaetes, polychaetes) and insects as size increased. Diet also varied with salinity, such that insects and zooplankton were important to YOY ( mm, n=611) in the limnetic-mesohaline Mill Creek area of upper Delaware Bay, while mysids were more important to YOY ( mm, n=97) in the mesohaline-polyhaline Dennis/Moores areas of lower Delaware Bay salt marshes (Nemerson 2001). Geographical variation in diet has also been noted within the Delaware Bay estuary (Nemerson 2001). Relative importance of individual prey items also varied intraannually in the Delaware River and the Hudson River estuaries (Miller 1963; Bath and O Connor 1985). Bath and O Connor (1985) found YOY (<110 mm) in the Hudson River ate amphipods, insect larvae, and cladocerans in the spring, while amphipods, isopods, and decapod shrimps were important in the summer. Relative importance of individual prey items also varied inter-annually in the lower Connecticut River (Marcy 1976a). Several other papers have been published on food habits of white perch, but they are not within the geographical range/age scope of this study, or were unable to be obtained (Hildebrand and Schroeder 1928; Smith 1947; Conover 1958; Thoits 1958; Elrod et al. 1981; Johnson 1983; Schaeffer and Margraf 1986; Parrish and Margraf 1994; Ruderhausen 1994; Klein-MacPhee 2002d; Moring and Mink 2002). Morone saxatilis, striped bass Several studies (n=8) have examined the food habits of YOY ( mm, n=2657) striped bass, Morone saxatilis, in Middle Atlantic Bight estuaries, from the Hudson River estuary, New York to Albemarle Sound, North Carolina (Tables 1, 2, 3). Important Prey items of YOY included a variety of invertebrates (polychaetes, insects, mysids, decapod shrimps, copepods, cladocerans, Limulus spp. eggs and larvae) and fish (Fundulus diaphanus, F. heteroclitus, Anchoa mitchilli, Gobiosoma bosc, Micropogonias undulatus). The diet of YOY shifted with ontogeny at approximately mm, from small invertebrates (mysids, insects, amphipods, copepods, cladocerans, polychaetes, isopods) at smaller sizes to larger invertebrates (decapod shrimps, crabs) and fish (Microgadus tomcod, Fundulus diaphanus, F. heteroclitus, Micropogonias undulatus, Anchoa mitchilli) at larger sizes (Markle and Grant 1970; Gardinier and Hoff 1982; Hartman and Brandt 1995; Cooper et al. 1998; Nemerson and Able 2003). The diet also varied with salinity. With salinity decreasing from mesohaline to limnetic/oligohaline portions of the estuaries, piscivory increased and mysid consumption decreased (Markle and Grant 1970; Boynton et al. 1981; Cooper et al. 1998; Nemerson and Able 2003). Also insects were more common food items in limnetic/oligohaline areas than in mesohaline areas (Markle and Grant 1970; Boynton et al. 1981). Geographical variation in diet has also been noted within the Potomac, Albemarle Sound and Delaware Bay estuaries (Boynton et al. 1981; Cooper et al. 1998, Nemerson and Able 2003) and between Virginia rivers (Markle and Grant 1970). Relative importance of individual prey items also varied both intra-annually in the Hudson River and Chesapeake Bay estuaries (Gardinier and Hoff 1982; Hartman and Brandt 1995) and inter-annually in the 19

26 Chesapeake Bay (Hurst and Conover 2001). Several other papers have been published on food habits of striped bass, but they are not within the geographical range/age scope of this study (Pearson 1938; McErlean 1961; Trent and Hassler 1966; Dovel 1968; Schaefer 1970; Holland and Yelverton 1973; Manooch 1973; Rulifson and McKenna 1987; Dew 1988; Robichaud-LeBlanc et al. 1997; Dunning et al. 1997; Bowman et al. 2000; Tupper and Able 2000; Klein-MacPhee 2002d; Walter et al. 2003; Harding and Mann 2003), or were unavailable (Curran and Ries 1937; Merriman 1941; Hollis 1952; Raney 1952; Bason 1971; Oviatt 1977; Setzler et al. 1980; Rudershausen 1994; Schulze 1996; Walter 1999; Griffin 2001; Nelson et al. 2003) Centropristis striata, black sea bass Family Serranidae Two studies examined the food habits of YOY ( mm, n=76) black sea bass, Centropristis striata, in Long Island Sound and Magothy Bay, Virginia (Tables 1 and 2). Important Prey of YOY included a variety of invertebrates (mysids, amphipods, crabs). Fish were a minor dietary component in Magothy Bay, Virginia (Kimmel 1973). At the same site, more than half of the diet of younger fish (30-91 mm, n=28) consisted of mysids, supplemented by amphipods, sediment, polychaetes, shrimps, isopods, cumaceans and plant remains, while larger fish ( mm, n=20) fed primarily on mud crab and commensal crabs, supplemented by mysids, amphipods, polychaetes, mollusks, shrimps, and sediment. Relative importance of individual prey items also varied intraannually in Magothy Bay, Virginia, such that during March and April more than half of the diet by volume was composed of mysids, while diet during spring and summer was more diverse, with mud crabs and amphipods consumed more frequently and mysids less frequently (Kimmel 1973). Several other papers have been published on food habits of black sea bass, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Bigelow and Schroeder 1953; Miller 1959; Cupka et al. 1973; Kendall 1977; Musick and Mercer 1977; Sedberry 1988; Hood et al. 1994; Bowman et al. 2000; Klein- MacPhee 2002g), or were unavailable (Link 1980). Pomatomus saltatrix, bluefish Family Pomatomidae Several studies (n=13) have examined the food habits of YOY ( mm FL, n=6853) bluefish, Pomatomus saltatrix, in Middle Atlantic Bight estuaries from Great South Bay, New York to lower Chesapeake Bay (Tables 1, 2, 3). Important Prey of YOY included a variety of fish (Menidia beryllina, M. menidia, Menidia spp., Brevoortia tyrannus, Alosa aestivalis, A. sapidissima, Alosa spp., Fundulus heteroclitus, F. majalis, Anchoa 20

27 mitchilli, Microgadus tomcod, Morone saxatilis, Morone spp.) and a few invertebrates (decapod shrimps, mysids). YOY exhibited cannibalistic behavior on a minor scale (Juanes et al. 1993; Buckel and Conover 1997; Gartland 2002). Two cohorts of YOY P.saltatrix, spring-spawned and summer-spawned, displayed differences in diet. Summer-spawned YOY exhibited a less diverse diet (almost exclusively fish, Anchoa mitchilli, with invertebrates rare) than spring-spawned YOY (fish, Morone saxatilis, Anchoa mitchilli, Alosa spp., Menidia menidia, supplemented by invertebrates, crab zoea and decapod shrimps) (Juanes et al. 1993; Juanes and Conover 1995; Buckel and Conover 1997; Buckel et al. 1999a). Several authors suggest the diet of YOY shifted with ontogeny at approximately mm FL, from a diet of fish supplemented by invertebrates, to a predominantly piscivorous diet (Grant 1962; Juanes et al. 1993; Juanes and Conover 1994; Juanes and Conover 1995; Able et al. 2003). Gartland (2002) suggested the shift to a piscivorous diet occurs much sooner, during estuarine recruitment (<33 mm FL). Geographical variation in diet has also been noted between estuaries (Juanes et al. 1994) and between estuarine and oceanic sites (Gartland 2002; Able et al. 2003). Relative importance of individual prey items also varied intra-annually (Juanes and Conover 1994; Hartman and Brandt 1995; Juanes and Conover 1995; Buckel and Conover 1997; Buckel et al. 1999a; Juanes et al. 2001; Gartland 2002). Gartland (2002) found the dominant prey of YOY ( mm, n=406) in Chesapeake Bay was Menidia menidia from late spring into early summer and then Anchoa mitchilli from mid-summer to fall. Relative importance of individual prey items also varied inter-annually (Friedland et al. 1988; Juanes et al. 1994; Juanes and Conover 1995; Buckel and Conover 1997; Buckel et al. 1999a). Juanes and Conover (1995) found decapod shrimps and fish of near equal importance (49.53%W, %W respectively) to spring-spawned YOY ( mm TL, n=176) in 1988, whereas in 1989 fish were much more important than decapod shrimps (67.95%W, 21.73%W) to the diet of YOY ( mm TL, n=534) in Great South Bay, New York. Several other papers have been published on food habits of bluefish, but they are not within the geographical range/age scope of this study (Breder 1922; Hildebrand and Schroeder 1928; Lux and Mahoney 1972; Wilk 1977; Lascara 1981; Naughton and Saloman 1984; Taylor 1987; Marks and Conover 1993; Creaser and Perkins 1994; Buckel et al. 1999b; Bowman et al. 2000; Harding and Mann 2001; Klein-MacPhee 2002a), or were unavailable (Lassiter 1962; Marks 1991; Juanes 1992; Buckel 1997). Caranx hippos, crevalle jack Family Carangidae One study examined the food habits of YOY ( mm TL, n=40) crevalle jack, Caranx hippos, in the Delaware River estuary (de Sylva et al. 1962) (Tables 1, 2, 3). YOY most frequently consumed mysid shrimp (Neomysis americana), decapod shrimps (Palaemonetes spp., Crangon spp.) and fishes (Gobiidae, Anchoa spp., Atherinidae). Several other papers have been published on food habits of crevalle jack, but they are not 21

28 within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Kwei 1978; Saloman and Naughton 1984; Klein-MacPhee 2002h). Lutjanus griseus, grey snapper Family Lutjanidae There have been no studies of the food habits of YOY (<110 mm TL) grey snapper, Lutjanus griseus, in Middle Atlantic Bight estuaries. One paper has been published on food habits, but it is not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928). Stenotomus chrysops, scup Family Sparidae One study examined the food habits of YOY ( mm, n=181) scup, Stenotomus chrysops, in Long Island Sound (Richards 1963) (Tables 1, 2). YOY concentrated on polychaetes, amphipods, other crustaceans and mollusks. The diet of YOY shifted with ontogeny at approximately mm, such that copepods and mysids were more prevalent in fish <100 mm, while larger fish consumed more mollusks, decapod shrimps and crabs. Several other papers have been published on food habits of scup, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Morse 1978; Sedberry 1983; Bowman and Michaels 1984; Bowman et al. 1987; Bowman et al. 2000; Klein-MacPhee 2002i), or were unavailable (Maurer and Bowman 1975; Michelman 1988). Family Sciaenidae Bairdiella chrysoura, silver perch Several studies (n=5) have examined the food habits of YOY (7-157 mm TL, n=630) silver perch, Bairdiella chrysoura, in Middle Atlantic Bight estuaries from the Delaware River to Beaufort, North Carolina (Tables 1, 2, 3). Important Prey of YOY included a variety of invertebrates (mysids, amphipods, decapod shrimps, polychaetes, oligochaetes, crabs, copepods) and fish (Anchoa mitchilli). The diet of YOY shifted with ontogeny at approximately mm, from a diet of small invertebrates (copepods, ostracods) to relatively larger invertebrates (decapod shrimp, crabs) and fish. In the York River estuary, Virginia YOY <40 mm SL fed mostly on copepods, but fed more on mysids, amphipods, and other crustaceans as they grew; fish became more Important Prey to individuals >70 mm SL (Chao and Musick 1977). At Beaufort, North Carolina, YOY (7-20 mm, n=30) fed chiefly on copepods, supplemented by ostracods, a few amphipods, cladocerans, an occasional mysid and chaetopods; YOY (25-50 mm, n=79) fed more frequently on relatively larger crustaceans (mysids, decapod shrimp, crabs, chaetopods, 22

29 mollusks) and more sparingly on copepods, ostracods and isopods, while YOY (50-80 mm, n=15) fed largely on mysids, shrimp, Gammarus spp., and chaetopods (Hildebrand and Cable 1930). Several other papers have been published on food habits of silver perch, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Reid 1954; Darnell 1958; Carr and Adams 1973; Kimmel 1973; Stickney et al. 1975; Adams 1976; Kleypas and Dean 1983; Bowman et al. 2000). Cynoscion regalis, weakfish Several studies (n=11) have examined the food habits of YOY ( mm, n=2989) weakfish, Cynoscion regalis, in Middle Atlantic Bight estuaries from Acushnet River, Massachusetts to Morehead City, North Carolina (Tables 1, 2, 3). Important Prey of YOY included a variety of invertebrates (mysids, decapod shrimps, amphipods, copepods, polychaetes, eggs) and fish (Anchoa mitchilli, Anchoa spp., Fundulus heteroclitus, Menidia menidia, Alosa aestivalis, Micropogonias undulatus). Also, YOY exhibit cannibalistic behavior (Thomas 1971; Hartman and Brandt 1995; Nemerson 2001). The diet of YOY shifted with ontogeny at approximately 60 mm, from a diet of small invertebrates (mysids, copepods) at smaller sizes to include more fish and decapod shrimps at larger sizes (Thomas 1971; Chao and Musick 1977; Goshorn 1990; Grecay 1990; Nemerson 2001). In the lower Delaware River, mysids and amphipods (68% frequency of occurrence (FO), 36% FO, respectively) were the most frequently occurring food items of YOY (<180 mm, n=494), copepods (12% FO) were generally restricted to the smallest size group (0-60 mm), while fishes (51% FO), and Crangon septemspinosa (17% FO) were more important to larger YOY ( mm) (Thomas 1971). Geographical variation in diet has also been noted within the Delaware River/Bay estuary (Thomas 1971; Grecay 1990; Nemerson 2001). In Delaware Bay marsh creeks, piscivory decreased with decreasing salinity, highest at meso-polyhaline lower bay sites Dennis Township/Moores Beach (45.5% weight [W]), intermediate at oligo-mesohaline Browns Run (20.4% W) and almost absent at limnetic-oligohaline Mill Creek (1.9% W). Mysid consumption was similar across Dennis/Moores, Browns Run and oligo-mesohaline Mad Horse Creek (26.0% W, 31.1% W, 33.6% W, respectively), but negligible at Mill Creek (0.6%W). Crustacean consumption was significantly higher at Browns Run (66.2% W) than at any of the other sites ( % W) (Nemerson 2001). Similarly mysids dominated the diet in all areas of Delaware Bay, but were of lesser importance in the upper bay, where benthic invertebrates and detritus were included in diet to greater extent than in lower and middle bay (Grecay 1990). Relative importance of individual prey items also varied seasonally in the Delaware Bay, where fish (35% FO) and decapod shrimps (15% FO) were more important in fall (September-October), especially Cynoscion regalis (23% FO) in September and decapod shrimps (Palaeonetes spp., Crangon spp.) (19.7% FO) in October (Thomas 1971). Diet also varied seasonally in Chesapeake Bay, such that Anchoa mitchilli (73.3 % W) was the main prey item supplemented by mysids (13.0% W) and grass shrimp (11.1% W) in summer (July- August). In fall (September-October), YOY consumed Anchoa mitchilli almost exclusively (97.9% W) and in winter (November-December) Anchoa mitchilli (58.2% W) 23

30 and Micropogonias undulatus (37.4% W) were the main prey items (Hartman and Brandt 1995). Several other papers have been published on food habits of weakfish, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Bigelow and Schroder 1953; Stickney et al. 1975; Wilk 1979; Lascara 1981; Mercer 1983; Taylor 1987; Lankford and Targett 1997; Bowman et al. 2000; Klein-MacPhee 2002j). Leiostomus xanthurus, spot Several studies (n=10) have examined the food habits of YOY (9-159 mm, n=6151) spot, Leiostomus xanthurus, in Middle Atlantic Bight estuaries from Delaware Bay to Cape Fear River, North Carolina (Tables 1, 2, 3). Important Prey of YOY included a variety of invertebrates (copepods, polychaetes, oligochaetes, amphipods, nematodes, bivalves, mysids, tanaids) and detritus. Sediment was also found frequently in stomachs. Fish (Family Atherinopsidae, Fundulus heteroclitus, Family Gobiidae, Micropogonias undulatus) were a minor dietary component (Table 3). The diet of YOY shifted with ontogeny at approximately 40 mm, from a diet of small invertebrates (copepods, mysids) when small to more benthic prey (polychaetes, oligochaetes, cumaceans, amphipods) when larger (Hildebrand and Cable 1930; Kimmel 1973; Hodson et al. 1981; Smith et al. 1984; O Neil and Weinstein 1987; Nemerson 2001). In Delaware Bay marsh creeks, mysid and meiobenthos consumption was highest by small individuals (20-40 mm); mysid consumption decreased to very low levels after 60 mm and meiobenthos consumption (mostly harpacticoid copepods) dropped steadily over the entire mm range, while annelid consumption increased from mm before leveling off thereafter (Nemerson 2001). Similarly, in the York River estuary, Virginia, small (16-20 mm) YOY consumed primarily copepods, whereas slightly larger individuals (26-30 mm) consumed more benthic prey (polychaetes, nematodes); larger YOY ( mm) consumed a wide variety of predominantly benthic organisms (polychaetes, amphipods, nematodes, oligochaetes), while the largest YOY examined (>100 mm) consumed primarily cumaceans and amphipods (O Neil and Weinstein 1987). Geographical variation in diet has also been noted within Delaware Bay (Nemerson 2001) such that annelids strongly dominated diet at meso-polyhaline Dennis Township (77.9% W), while only 23.9% W and 31.5% W at oligo-mesohaline Browns Run and Mad Horse Creek, respectively. Meibenthos were common prey, but significantly higher at Browns Run (29.2% W) than at Dennis and Mad Horse (12.4% W at each), while crustacean consumption was significantly higher at Browns Run (19.6%) and Mad Horse (19.1%) than at Dennis (3.3%W). Spatial diet variation was also noted at York River, Virginia, where dominant items consumed by YOY at meso-oligohaline, Goalder s Creek included nereid polychaetes, clam siphons, gammarid amphipods and harpacticoid copepods, while YOY from polyhaline, Blevins Creek consumed more nematodes, maldanid polychaetes and oligochaetes (O Neil and Weinstein 1987). Relative importance of individual prey items also varied intra-annually (Hines et al. 1990; Nemerson 2001). In the Rhode River subestuary of Chesapeake Bay, YOY ( mm, n=187) consumed 24

31 mainly amphipods and polychaetes early in the season, primarily clam (Macoma balthica), siphons, small crustaceans and meiofauna in June and a variety of polychaetes throughout the year (Hines et al. 1990). Several other papers have been published on food habits of spot, but they are not within the geographical range/age scope of this study (Welsh and Breder 1923; Hildebrand and Schroeder 1928; Pearson 1929; Gunter 1945; Reid 1954; Darnell 1958; Dawson 1958; Springer and Woodburn 1960; Thomas 1971; Parker 1971; Stickney and McGeachin 1975; Peters and Kjelson 1975; Stickney et al. 1975; Adams 1976; Chao and Musick 1977; Kobylinski and Sheridan 1979; Govoni et al. 1983; Currin et al. 1984; Smith and Coull 1987; Archambault and Feller 1991; Miltner et al. 1995; Bowman et al. 2000; Klein-MacPhee 2002j) or were unavailable (Smith 1907; Van Engle and Joseph 1968). Menticirrhus saxatilis, northern kingfish Two studies examined the food habits of YOY ( mm, n=139) northern kingfish, Menticirrhus saxatilis, in the Delaware River estuary and York River, Virginia (Tables 1, 2, 3). Important Prey of YOY included a variety of invertebrates (polychaetes, mysid shrimps, amphipods, copepods), fish (Family Clupeidae) and detritus. Several other papers have been published on food habits of northern kingfish, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Gunter 1945; Springer and Woodburn 1960; Irwin 1970; Thomas 1971; Kimmel 1973; McMichael and Ross 1987; Bowman et al. 2000; Klein-MacPhee 2002j). Micropogonias undulatus, Atlantic croaker Several studies (n=5) have examined the food habits of YOY ( mm, n=2273) Atlantic croaker, Micropogonias undulatus, in Middle Atlantic Bight estuaries from Delaware Bay to North Carolina (Tables 1, 2). Important Prey of YOY included a variety of invertebrates (copepods, polychaetes, mysids, amphipods, oligochaetes, gastropods, bivalves, decapod shrimps, ostracods) and fish. The diet of YOY shifted with ontogeny in Delaware Bay (Thomas 1971) and Delaware Bay marsh creeks (Nemerson 2001), such that YOY consumed more fish, crustaceans and annelids and fewer mysids and zooplankton with growth. Spatial variation in diet has also been noted between marsh creeks within Delaware Bay such that annelids and crustaceans were Important Prey at all sites. Piscivory was highest at meso-polyhaline lower bay sites Dennis/Moores (9.2% W). Mysids were more important at Dennis/Moores and oligomesohaline Mad Horse Creek (22.0% W and 23.8% W, respectively), while zooplankton and insects were important (19.8% W, 15.1% W) at limnetic-oligohaline Mill Creek (Nemerson 2001). Relative importance of individual prey items also varied intraannually in Delaware Bay marsh creeks, such that amphipods became more important, while copepods and decapod shrimps declined in importance from September through November (Nemerson 2001). 25

32 Several other papers have been published on food habits of Atlantic croaker, but they are not within the geographical range/age scope of this study (Pearson 1929; Hildebrand and Cable 1930; Gunter 1945; Reid 1955; Darnell 1958; Hansen 1969; Kimmel 1973; Stickney et al. 1975; Stickney and McGeachin 1975; Sheridan 1979; Kobylinski and Sheridan 1979; Govoni et al. 1983; Currin et al. 1984; Bowman et al. 2000). Pogonias cromis, black drum Two studies examined the food habits of YOY ( mm, n=268) black drum, Pogonias cromis, in the Delaware River estuary (Tables 1, 2, 3). Important Prey of YOY included a variety of invertebrates (polychaetes, oligochaetes, bivalves, insects, amphipods). Fish (Menidia spp., Family Clupeidae, Anchoa mitchilli) were a minor dietary component (Table 3). Geographical variation in diet has been noted within the lower Delaware Bay estuary, such that amphipods were of greatest numerical importance to YOY in both the river and creek locations, while annelids were of secondary individuals in the river and copepods were of secondary importance in creeks (Thomas 1971). Several other papers have been published on food habits of black drum, but they are not within the geographical range/age scope of this study (Pearson 1929; Simmons and Breuer 1962; Music and Pafford 1984; Peters and McMichael 1990; Klein-MacPhee 2002j). Chaetodon ocellatus, spotfin butterflyfish Family Chaetodontidae There have been no studies of the food habits of YOY (<63 mm) spotfin butterflyfish, Chaetodon ocellatus, in Middle Atlantic Bight estuaries. However, one other paper has been published on food habits, but they are not within the geographical range/age scope of this study (Pitts 1991; McBride 2002). Mugil cephalus, striped mullet Family Mugilidae There have been no studies of the food habits of YOY (<200) striped mullet, Mugil cephalus, in Middle Atlantic Bight estuaries However, several other papers have been published on food habits, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Hiatt 1947a,b; Ebeling 1957; Anderson 1957; Darnell 1958; Arnold and Thompson 1958; Wood 1964; Suzuki 1965; Odum 1966; Odum 1968; Odum 1970; Odum 1971; Odum and Heald 1972; Kimmel 1973; De Silva and Wijeyaratne 1977; Blaber and Whitfield 1977; Marais 1980; Loftus et al. 1983; Eggold and Motta 1992; Thompson 2002). 26

33 Mugil curema, white mullet There have been no studies of the food habits of YOY (<200 mm) white mullet, Mugil curema, in Middle Atlantic Bight estuaries However, several other papers have been published on food habits, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Kimmel 1973; Moore 1974; Gonzalez and Alvarez-Lajonchere 1978; DeLancey 1989; Thompson 2002). Sphyraena borealis, northern sennet Family Sphyraenidae One study examined the food habits of YOY ( mm, n=85) northern sennet, Sphyreana borealis, in the Delaware River estuary (de Sylva et al. 1962) (Tables 1, 2, 3). YOY primarily consumed fishes, Menidia spp., most frequently, as well as Alosa spp., Anchoa spp., and Gobiosoma spp. with decreasing frequency. YOY also exhibit cannibalistic behavior on a minor scale. Mysids were also Important Prey. Several other papers have been published on food habits of northern sennet, but they are not within the geographical range/age scope of this study (Rountree 2002), or were unavailable (Smith 1907; Sumner et al. 1913). Tautoga onitis, tautog Family Labridae One study examined the food habits of YOY (31-71 mm, n=36) tautog, Tautoga onitis, in Great South Bay, New York (Grover 1982) (Tables 1, 2). YOY consumed primarily copepods (45.7% composition) and amphipods. Several other papers have been published on food habits of tautog, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Richards 1963; Olla et al. 1974; Olla et al. 1975; Steimle and Ogren 1982; Lindquist et al. 1985; Bowman et al. 2000; Munroe 2002b; Able and Duffy-Anderson 2005), or were unavailable (Dorf 1994). Tautogolabrus adspersus, cunner One study examined the food habits of YOY (11-64 mm, n=54) cunner, Tautogolabrus adspersus, in Great South Bay, New York (Grover 1982) (Tables 1, 2). YOY consumed predominantly copepods and amphipods. Fish were a minor dietary component. Dependence on copepods decreased with growth, such that copepods comprised almost the entire diet (97.2%) at 15.5 mm, whereas at 45.9 mm, copepods comprised only 32.1% of the diet. 27

34 Several other papers have been published on food habits of cunner, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Bigelow and Schroeder 1953; Richards 1963; Green and Farwell 1971; Chao 1973; Shumway and Stickney 1975; Olla et al. 1975; Edwards et al. 1982; Whoriskey 1983; Green et al. 1984; Green et al. 1985; Harris 1986; Bowman et al. 2000; Munroe 2002b), or were unavailable (Sand 1982). Pholis gunnellus, rock gunnel Family Pholidae There have been no studies of the food habits of YOY (<73 mm) rock gunnel, Pholis gunnellus, in Middle Atlantic Bight estuaries. However, several other papers have been published on food habits, but they are not within the geographical range/age scope of this study (Stroud 1939; Backus 1957; Sawyer 1967; Proudfoot 1975; Paperno 1983; Cheetham and Fives 1990; Collette 2002b), or were unavailable (Qasim 1957). Astroscopus guttatus, northern stargazer Family Uranoscopidae There have been no studies of the food habits of YOY (<90 mm) northern stargazer, Astroscopus guttatus, in Middle Atlantic Bight estuaries. One other paper has been published on food habits, but it was not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928). Family Blenniidae Hypsoblennius hentz, feather blenny There have been no studies of the food habits of YOY (<90 mm) feather blenny, Hypsoblennius hentz, in Middle Atlantic Bight estuaries Several other papers have been published on food habits, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Springer and Woodburn 1960; Kimmel 1973). Family Ammodytidae Ammodytes americanus, American sand lance Several studies (n=4) have examined the food habits of YOY (8-159 mm, n=740) American sand lance, Ammodytes americanus, in Long Island Sound and Great South Bay, New York (Tables 1, 2, 3). Important Prey of YOY included a variety of invertebrates (copepods, crabs, cirripedes, bivalves, gastropods, cnidarians, mysid shrimps, decapod shrimps, cladocerans) and phytoplankton. Fish were a minor dietary component. YOY exhibit cannibalistic behavior (Richards 1963). In Long Island Sound, the diet of larvae (<8-24 mm, n=175) shifted from phytoplankton to various stages of 28

35 copepods with growth. For example, small larvae (<8 mm) contained only phytoplankton, while larger larvae ( mm) consumed copepod nauplii. At larger sizes ( mm), the larvae consumed both nauplii and copepodites, and at larger sizes ( mm) they consumed greater amounts of copepodites and Acartia hudsonica adults than nauplii. The largest larvae examined ( 24 mm) consumed A. hudsonica adults almost exclusively. Relative importance of individual prey items also varied intra and inter-annually in Long Island Sound (McKown 1984), such that YOY examined from June 1982 contained large quantities of crab zoea, while those examined from June 1983 contained mostly cladocerans and snail veligers. Acartia spp. copepodites and adults comprised 60% and 10%, respectively, of the diet of YOY examined from July 1983, as well as small percentages of Temora adults and copepodites, harpacticoid copepods and crab zoea. Adult Labidocera and adult Acartia spp. comprised more than 30% and 20%, respectively, of the diet of individuals collected on December Several other papers have been published on food habits of American sand lance, but they are not within the geographical range/age scope of this study (Meyer et al. 1979; Sedberry 1983; Bowman and Michaels 1984; Bowman et al. 1987; Scott and Scott 1988; Fortier et al. 1995; Nizinski 2002; Zamon 2003), or were unavailable (Covill 1959). Gobionellus boleosoma, darter goby Family Gobiidae There have been no studies of the food habits of YOY (<40 mm) darter goby, Gobionellus boleosoma, in Middle Atlantic Bight estuaries. However, several other papers have been published on food habits of darter goby, but they are not within the geographical range/age scope of this study (Carle and Hastings 1982; Fitzhugh and Fleeger 1985; Toepfer and Fleeger 1995; Gregg and Fleeger 1997). Gobiosoma bosc, naked goby There have been no studies of the food habits of YOY (<60 mm) naked goby, Gobiosoma bosc, in Middle Atlantic Bight estuaries. One other paper has been published on food habits of naked goby, but it is not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928). Gobiosoma ginsburgi, seaboard goby One study examined the food habits of YOY (18-42 mm SL, n=49) seaboard goby, Gobiosoma ginsburgi, in Sakonnet River, Rhode Island (Munroe and Lotspeich 1979) (Tables 1, 2). Crustaceans were the most frequent and abundant food items, predominantly harpacticoid copepods (60%) supplemented by gammaridean amphipods (18%), polychaetes (13%) and mollusks (11%). 29

36 Several other papers have been published on food habits of seaboard goby, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Richards 1963; Kimmel 1973). Peprilus triacanthus, butterfish Family Stromateidae One study examined the food habits of YOY ( mm, n=19) butterfish, Peprilus triacanthus, in Long Island Sound (Tables 1, 2). Available crustaceans were the majority of YOY prey, most frequently Acartia spp. copepods. Several other papers have been published on food habits of butterfish, but they are not within the geographical range/age scope of this study (Fritz 1965; Horn 1970; Oviatt and Kremer 1977; Bowman and Michaels 1984; Bowman et al. 1987; Bowman et al. 2000; Klein-MacPhee 2002k). Scophthalmus aquosus, windowpane Family Scophthalmidae Several studies (n=3) have examined the food habits of YOY ( mm, n=520) windowpane, Scophthalmus aquosus, in Middle Atlantic Bight estuaries from Long Island Sound to Delaware River (Tables 1, 2). Important Prey of YOY included a variety of invertebrates (mysids, copepods, amphipods, decapod shrimps, gastropods). Fish were a minor dietary component. The diet of YOY in the Great Bay-Little Egg estuary, New Jersey shifted with increasing size (Haberland 2002), such that larvae (<5 mm, n=103) consumed large amounts of gastropod veligers and calanoid copepods, small YOY (6-40 mm, n=173) consumed a mix of Neomysis americana, copepods and amphipods, larger YOY (40-70 mm, n=69) consumed large amounts of N. americana along with increasing amounts of Crangon, and the largest YOY examined (70-90 mm, n=38) consumed a large increase in the amount of Crangon, increasing amounts of Gammarus spp. and a reduction in the amount of N. americana. Several other papers have been published on food habits of windowpane, but they are not within the geographical range/age scope of this study (Bigelow and Schroeder 1953; Kimmel 1973; Moore 1974; Hacunda 1981; Langton and Bowman 1980; Bowman et al. 2000; Klein-MacPhee 2002l). Etropus microstomus, smallmouth flounder Family Paralichthyidae There have been no studies of the food habits of YOY (<120 mm) smallmouth flounder, Etropus microstomus, in Middle Atlantic Bight estuaries However, one other paper has 30

37 been published on food habits of smallmouth flounder, but was not within the geographical range/age scope of this study (Richards 1963). Paralichthys dentatus, summer flounder Several studies (n=9) have examined the food habits of YOY ( mm, n=1211) summer flounder, Paralichthys dentatus, in Middle Atlantic Bight estuaries from Great Bay-Little Egg Harbor, New Jersey to Pamlico Sound, North Carolina (Tables 1, 2, 3). Important Prey of YOY included a variety of invertebrates (mysid shrimps, decapod shrimps, polychaetes, copepods, crabs) and fish (Menidia menidia, Fundulus heteroclitus, Leiostomus xanthurus). Detritus and sediment were also prominent. The diet of small YOY (9-20 mm, n=173) shifted with growth during the immigration period (January- March, 1988) to the Newport and North Rivers of North Carolina (Burke 1995), such that the smallest individuals (9-12 mm, n=110) consumed mainly a mixture of polychaetes, polychaete tentacles, mysids and harpactacoid copepods. With growth (13-16 mm, n=53), polychaete tentacles decreased in importance, harpacticoids were no longer consumed, and polychaetes and calanoid copepods became more important. With further growth (17-20 mm, n=10), mysids became Important Prey in addition to polychaetes, supplemented by calanoid copepods and amphipods. In Pamlico Sound, North Carolina, the diet shifted such that, smaller ( mm, n=470) individuals consumed predominantly mysids (42% Volume) and fishes (38%V), while larger YOY ( mm, n=49) consumed more fish (56%V) and decapod shrimps (36%V) (Powell and Schwartz 1979). Some changes in feeding patterns related to temporal abundance of a food item in a discrete area were also noted in Pamlico Sound, such that during summer a relatively large amount of amphipods were consumed by fish at one collecting locality, while during winter and spring shrimp were relatively common near inlet stations. Several other papers have been published on food habits of summer flounder, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Poole 1961; Poole 1964; Smith 1969; Adams 1976; Smith and Daiber 1977; Taylor 1987; Timmons 1995; Bowman et al. 2000; Klein-MacPhee 2002m). Family Pleuronectidae Pseudopleuronectes americanus, winter flounder Several studies (n=10) have examined the food habits of YOY (3-300 mm, n=2681) winter flounder, Pseudopleuronectes americanus, in Middle Atlantic Bight estuaries from the lower Pettaquamscutt River, Rhode Island to Rehoboth Bay, Delaware (Tables 1, 2, 3). Important Prey of YOY included a variety of invertebrates (polychaetes, amphipods, copepods, isopods, bivalves, ostracods, gastropods, nemertines, decapod shrimps, oligochaetes, mysid shrimps, cumaceans, crabs) and detritus. Fish (Menidia menidia, Anchoa mitchilli, Gobiosoma bosc) were a minor dietary component. The diet of YOY shifted with ontogeny at approximately mm, from a diet of small polychaetes, copepods and invertebrate eggs when smaller to larger polychaetes, 31

38 amphipods and decapod shrimps when larger (Mulkana 1964; Stehlik and Meise 2000; Vivian et al. 2000). The diet of larvae (3-8 mm, n=140) shifted in the Mystic River estuary, Connecticut (Pearcy 1962) such that, the smallest larvae (3-5 mm, n=37) consumed copepod nauplii most commonly, invertebrate eggs and larval polychaetes with decreasing frequency, while phytoplankton (pinnate and filamentous diatoms) were noted occasionally in specimens (3-4 mm). Small polychaetes were the major identifiable food of slightly larger larvae (6-8 mm, n=25), supplemented by nauplii, harpactacoid and calanoid copepods. YOY ( mm, n=1291) in the Hudson-Raritan estuary, New York/New Jersey also showed an ontogenetic diet shift (Stehlik and Meise 2000). Copepods and Spionidae polychaetes were prominent in diet of small (15-49 mm) individuals and decreased in importance as size increased. At 50 mm, there was a sharp reduction in the importance of copepods in the diet. Amphipods were the most consistent prey in the much broader diet of larger ( mm) individuals, supplemented by the bivalve, Mya arenaria (whole and siphons), the gastropod, Nassarius spp., various polychaetes, isopods, amphipods, mysid shrimps and the decapod shrimp, Crangon septemspinosa. Vivian et al. (2000) found similar results in the Hudson River estuary. YOY (10-80 mm, n=123) also showed ontogenetic diet shifts in the lower Pettaquamscutt River estuary, Rhode Island at approximately 60 mm, such that polychaetes, amphipods and Crangon increased in frequency with increasing size, while copepods, ostracods, and invertebrate eggs decreased in frequency with size (Mulkana 1964). Several other papers have been published on food habits of winter flounder, but they are not within the geographical range/age scope of this study (Hildebrand and Schroeder 1928; Kennedy and Steele 1971; Wells et al. 1973; Frame 1974; Langton and Bowman 1980; Hacunda 1981; Worobec 1984; Frank and Leggett 1984; MacDonald and Green 1986; MacDonald and Waiwood 1987; Grimes et al. 1989; Franz and Tanacredi 1992; Steimle et al. 1994; Timmons 1995; Stoner et al. 1999; Bowman et al. 2000; Manderson et al. 2000; Duffy-Anderson and Able 2001; Klein-MacPhee 2002n; Able and Duffy- Anderson 2005), or were unavailable (Linton 1921). Trinectes maculatus, hogchoker Family Soleidae One study examined the food habits of YOY (21-60 mm, n=83) hogchoker, Trinectes maculatus, in the York River, Virginia (Smith et al. 1984) (Tables 1, 2). YOY consumed predominantly amphipods, supplemented by detritus, polychaetes, copepods, and clam siphons. Several other papers have been published on food habits of hogchoker, but they are not within the geographical range/age scope of this study (Castagna 1955; Darnell 1958; Springer and Woodburn 1960; de Sylva et al. 1962; Koski 1973; Carr and Adams 1973; Kimmel 1973; Hines et al. 1990; Klein-MacPhee 2002o), or were unavailable (Van Engle and Joseph 1968; Schwartz 1997b). 32

39 Sphoeroides maculatus, northern puffer Family Tetraodontidae One study examined the food habits of YOY ( mm, n=22) northern puffer, Sphoeroides maculatus, in Long Island Sound (Tables 1, 2). YOY consumed predominantly polychaetes, crustaceans, most importantly amphipods, and pelecypod mollusks. Several other papers have been published on food habits of northern puffer, but they are not within the geographical range/age scope of this study (Welsh and Breder 1922; Hildebrand and Schroeder 1928; Randall 1967; Shipp and Yerger 1969; Klein-MacPhee 2002p), or were unavailable (Isaacson 1963). Limitations of This Study DISCUSSION The accuracy of our interpretation of food habits for young-of-the-year fishes is based entirely on the available literature. Thus, the ability to interpret food habits accurately is based on a variety of characteristics of each study including sample fish size, size range, method of sample collection, and frequency of examination (i.e. over days, months, years). Further, the breadth of the studies for each species can be influenced by the geographical extent of sample collection, estuarine habitat and especially salinity because it may influence prey availability, and those characteristics listed above. Perhaps most importantly, the measures of prey importance we have used are relatively crude because they are assigned to just three levels of importance (important, minor and absent). This was necessary because the techniques employed by individual studies varied widely and thus were not easily comparable. Another concern is that the selection of prey categories may have influenced the interpretation of prey importance because some categories are overlapping (e.g. copepods and zooplankton). The selection of prey categories was largely based on the taxa provided in the available literature. Many of these concerns have been expressed in other studies of food habits of fishes (Garrison and Link, 2000; Stouder et al. 1994; Ross, 1986; Livingston, 1982; Helfman, 1978; Darnell, 1961; Hynes, 1950). Conclusions The importance of copepods, amphipods, mysids, decapod shrimp and crabs, polychaetes, insects/arachnids, as found in this study, is typical for young-of-the-year in other estuarine studies in other regions (Festa, 1979, demorais and Bodiou 1984, Fitzhugh and Fleeger 1985, Gee 1989, Coull 1990, Feller et al. 1990, Coull et al. 1995). The importance of small prey, as indicated by the dominance of the above groups probably simply reflects morphological influence of gap limitations (Karpouzi and 33

40 Stergiou, 2003) of the small, young-of-the-year fishes that occurs across all taxa regardless of habitat. The wide variability in prey types within a species is likely due to prey availability that varies along the salinity gradient in estuaries in general. The importance of fish as prey is somewhat surprising given the often repeated theme that estuaries serve as refugia from predation (Joseph 1972, Weinstein 1979, Boesch and Turner 1984, Blaber 2001). Most of the dominant prey species are, not unexpectantly, small species that are abundant in many estuarine habitats (Fundulus heteroclitus, Menidia menidia, and Anchoa mitchilli [Table 2]). Others, such as Morone saxatilis, were reported in the diet of species that become piscivorous at relatively small sizes (Pomotomus saltatrix) and also reflect the many studies in the Hudson River where M. saxatilis is abundant, (see below). ACKNOWLEDGMENTS We would like to thank Sue Viscardi and Bobbie Zlotnik for editorial assistance. This is report was funded by support from the Rutgers University Marine Field Station. This is Institute of Marine and Coastal Sciences Contribution # LITERATURE CITED Able, K.W Killifishes. Order Cyprinodontiformes. Pp In: Collette, B.B., and G. Klein-MacPhee (eds.), Bigelow and Schroeder's Fishes of the Gulf of rd Maine, Smithsonian Institution, 3 edition. Able, K.W. and J.T. Duffy-Anderson Impacts of piers on juvenile fishes and selected invertebrates in the lower Hudson River. In: The Hudson River, J. Levinton and C. Hiles (eds.), Cambridge University Press, Cambridge, England. Able, K.W. and M.P. Fahay The First Year in the Life of Estuarine Fishes in the Middle Atlantic Bight. Rutgers University Press, New Brunswick, NJ. 342 pp. Able, K.W., P. Rowe, M. Burlas, and D. Byrne Use of ocean and estuarine habitats by young-of-year bluefish (Pomatomus saltatrix) in the New York Bight. Fish. Bull., U.S. 101: Adams, S.M Feeding ecology of eelgrass fish communities. Trans. Amer. Fish. Soc. 105(4): Alexander, L.C Feeding chronology and food habits of the tomcod Microgadus tomcod (Walbaum) and winter flounder Pseudopleuronectes americanus Walbaum in Montsweag Bay (Sheepscot River), Maine. M.S. thesis, Univ. of Maine, Orono. 36 pp. 34

41 Allen, D.M., W.S. Johnson, and V. Ogburn-Matthews Trophic relationships and seasonal utilization of salt-marsh creeks by zooplanktivorous fishes. Envir. Biol. Fish. 42: Allen, E.A., P.E. Fell, M.A. Peck, J.A. Gieg, C.R. Guthke, and M.D. Newkirk Gut contents of common mummichogs, Fundulus heteroclitus L., in a restored impounded marsh and in natural reference marshes. Estuaries 17(2): Anderson, W.W Early development, spawning, growth, and occurrence of the silver mullet (Mugil curema) along the south Atlantic coast of the United States. Fish. Bull., U.S. 119: Archambault, J.A., and R.J. Feller Diel variation in gut fullness of juvenile spot, Leiostomus xanthurus. Estuaries 14(1): Arnold, E.L., and J.R. Thompson Offshore spawning of the striped mullet, Mugil cephalus, in the Gulf of Mexico. Copeia 1958(2): Atkinson, C.E Feeding habits of adult shad (Alosa sapidissima) in fresh water. Ecology 32(3): Auth, T.D Interannual and regional patterns of abundance, growth, and feeding ecology of larval bay anchovy (Anchoa mitchilli) in Chesapeake Bay. M.S. thesis, University of Maryland, College Park. Backus, R.H The fishes of Labrador. Bull. Am. Mus. Nat. Hist. 113: Baker-Dittus, A.M Foraging patterns of three sympiatric killifish. Copeia 1978(3): Barans, C.A Distribution, growth and behavior of the spotted hake in the Chesapeake Bight. M.A. thesis, College of William and Mary, Virginia. 54 pp. Bason, W.H Ecology and early life history of striped bass, Morone saxatilis, in the Delaware Estuary. M.S. thesis, Cornell Univ. 122 pp. Bath, D.W., and J.M. O'Conner Food preferences of white perch in the Hudson River estuary. N.Y. Fish Game J. 32(1): Battle, H.I., A.C. Huntsman, A.M. Jeffers, G.W. Jeffers, W.H. Johnson, and N.H. McNairn Fatness, digestion, and food of Passamoquoddy young herring. J. Biol. Bd. Can. 2: Bayliff, W.H The life history of the Atlantic silverside, Menidia menidia (L.). Chesapeake Biol. Lab. Publ. No pp. 35

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43 Boynton, W.R., T.T. Polgar, and H.H. Zion Importance of juvenile striped bass food habits in the Potomac Estuary. Trans. Amer. Fish. Soc.110: Branstetter, S Smooth dogfishes. Family Triakidae. In: Collette, B.B., and G. Klein-MacPhee (eds.), Bigelow and Schroeder's Fishes of the Gulf of Maine, Smithsonian Institution, 3rd. Edition pp. Breder, C.M., Jr The food of Mustelus canis (Mitchill) in mid-summer. Copeia 1921(101): Breder, G.C Observations on young bluefish. Copeia 1922(106): Brinley, F.J., and R.E. Bowen Some feeding habits of the common eel, (Anguilla bostoniensis). Copeia 1935(3):140. Bromley, P.J., T. Watson, and J.R.G. Hislop Diel feeding patterns and development of food webs in pelagic 0-group cod (Gadus morhua), haddock (Melanogrammus aeglefinus), whiting (Merlangius merlangus), saithe (Pollachius virens), and Norwary pout (Trisopterus esmarkii) in northern North Sea. ICES Journal of Marine Science 54: Buckel, J.A Impact of bluefish (Pomatomus saltatrix) predation on estuarine and continental shelf fishes. Ph.D. dissertation, State University of New York, Stony Brook. Buckel, J.A. and D.O. Conover Movements, feeding periods, and daily ration of piscivorous young-of-the-year bluefish, Pomatomus saltatrix, in the Hudson River estuary. Fish. Bull., U.S. 95: Buckel, J.A., M.J. Fogarty, and D.O. Conover. 1999a. Foraging habits of bluefish Pomatomus saltatrix, on the U.S. east coast continental shelf. Fish. Bull., U.S. 97: Buckel, J.A., D.O. Conover, N.D. Steinberg, and K.A. McKown. 1999b. 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). Can. J. Fish. Aquat. Sci. 56: Burbidge, R.G Distribution, growth, selective feeding, and energy transformation of young-of-the-year blueback herring, Alosa aestivalis (Mitchill) in the James River, Virginia. Trans. Am. Fish. Soc. 103: Burke, J.S Role of feeding and prey distribution of summer and southern flounder in selection of estuarine nursery habitats. J. Fish Biol. 47:

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65 Odum, W.E., and E.J. Heald Trophic analyses of an estuarine mangrove community. Bull. Mar. Sci. 22(3): Ogden, J.C Relative abundance, food habits, and age of the American eel, Anguilla rostrata (LeSueur), in certain New Jersey streams. Trans. Amer. Fish. Soc. 1970(1): O'Gorman, R Predation by rainbow smelt (Osmerus mordax) on young-of-theyear alewives (Alosa pseudoharengus) in the Great Lakes. Prog. Fish Cult. 36: Ojeda, F.P., and J.H. Dearborn Feeding ecology of benthic mobile predators: experimental analyses of their influence in rocky subtidal communities of the Gulf of Maine. J. Exp. Mar. Biol. Ecol. 149: Olla, B.L., A.J. Bejda, and A.D. Martin Daily activity, movements, feeding, and seasonal occurrence in the tautog, Tautoga onitis. Fish. Bull., U.S. 72(1): Olla, B.L., A.J. Bejda, and A.D. Martin Activity, movements, and feeding behavior of the cunner, Tautogolabrus adspersus, and comparison of food habits with young tautog, Tautoga onitis, off Long Island, New York. Fish. Bull., U.S.73(4): O'Neil, S.P, and M.P. Weinstein Feeding habits of spot, Leiostomus xanthurus, in polyhaline verso meso-oligohaline tidal creeks and shoals. Fish. Bull., U.S. 85(4): Oviatt, C.A Menhaden, sport fish, and fisherman. Univ. of Rhode Island Marine Technical Report No pp. Oviatt, C.A., and P.M. Kremer Predation on the ctenophore, Mnemiopsis leidyi, by butterfish, Peprilus triacanthus, in Narragansett Bay, Rhode Island. Chesapeake Sci. 18(2): Paperno, R Geographic variation in the rock gunnel (Pisces: Pholidae) Pholis gunnellus (L.). M.S. thesis, Adelphi Univ. 57 pp. Parker, J.C The biology of the spot, Leiostomus xanthurus Lacepede, and Atlantic croaker, Micropoqon undulatus in two Gulf of Mexico nursery areas. Tex. AandM Univ. Sea Grant Coll. TAMU-SG pp. Parrish, D.L., and F.J. Margraf Spatial and temporal patterns of food use by white perch and yellow perch in Lake Erie. J. Freshwater Ecol. 9(1):

66 Pearcy, W.G Ecology of an estuarine population of winter flounder, Pseudopleuronectes americanus (Walbaum). Parts I-IV. Bull. Bingham Oceanogr. Coll. 18(1). Pearson, J.C Natural history and conservation of the redfish and other commercial Sciaenids on the Texas coast. Bull. Bur. Fish. XLIV: Pearson, J.C The life history of the striped bass, or rockfish, Roccus saxatilis (Walbaum). Fish. Bull., U.S. 49: Peck, J.I On the food of the menhaden. Bull. U.S. Fish Com. 13: Peters, D.S., and M.A. Kjelson Consumption and utilization of food by various postlarval and juvenile fishes of North Carolina estuaries. In: L.E. Cronin (ed.) Estuarine Research, Vol.1. Academic Press, New York pp. Peters, K.M., and R.H. McMichael, Jr Early life history of the black drum Pogonias cromis (PISCES: SCIAENIDAE) in Tampa Bay, Florida. Northeast Gulf Sci. 11(1): Petrillo, A.P Trophic structure of a tidal marsh finfish community (Branford, Connecticut). M.S. thesis, Southern Connecticut State University, New Haven. 101 pp. Phelps, G.B Ontogenetic variation in the diet of the inshore lizardfish, Synodus foetens. Dept. Biological Sciences, Florida Tech., Melbourne, FL. Pitts, P.A Comparative use of food and space by three Bahamian butterflyfishes. Bull. Mar. Sci. 48: Polgar, T.T Larval retention: Transport and behavior, on differential mortality. Estuaries 4:276. Poole, J.C Age and growth of the fluke in Great South Bay and their significance to the sport fishery. N.Y. Fish Game J. 8:1-18. Poole, J Feeding habits of the summer flounder in Great South Bay. N.Y. Fish Game J. 11: Poulin, R., and G.J. FitzGerald Early life histories of three sympatric sticklebacks in a salt-marsh. J. Fish Biol. 34: Powell, A.B., and F.J. Schwartz Food of Paralichthys dentatus and P. lethostigma (Pisces: Bothidae) in North Carolina estuaries. Estuaries 2(4):

67 Price, J.W A study of the food habits of some Lake Erie fish. Bull. Ohio Biol. Surv. New Ser. 2. Proudfoot, L.A The biology of the rock gunnel Pholis gunnellus (Linnaeus). B.S. Hon. thesis, Memorial Univ., St. John's, Newfoundland. 40 pp. Pyle, A.B Some aspects of the life history of Cyprinodon variegatus Lacepede 1803, in New Jersey and its reaction to environmental changes. Master s thesis, Rutgers, The State University of New Jersey, New Brunswick. Qasim, S.Z The biology of Centronotus gunnellus (L.) (Teleostei). J. Anim. Ecol. 26: Randall, J.E Food habits of reef fishes of the West Indies. Univ. Miami Trop. Oceanogr. Stud. No Raney, E.C The life history of the striped bass, Roccus saxatilis (Walbaum). Bull. Bingham Oceanogr. Coll. 14(1):5-97. Reid, G.K., Jr An ecological study of the Gulf of Mexico fishes, in the vicinity of Cedar Key, Florida. Bull. Mar. Sci. Gulf Carib. 4:1-94. Reid, G.K., Jr A summer study of the biology and ecology of East Bay, Texas. Part II. The fish fauna of East Bay, the Gulf Beach, and summary. Texas J. Sci. 4: Richards, S.W The demersal fish population of Long Island Sound. Bull. Bingham Oceanogr. Coll. 18(2). Richards, S.W., J.M. Mann, and J.A. Walker 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(4): Roberts, S.C Biological and fisheries data on northern searobin, Prionotus carolinus (Linnaeus). NMFS Sandy Hook Lab., Tech. Ser. Rept. No. 13. Roberts-Goodwin, S.C Biological and fisheries data on striped searobin, Prionotus evolans (Linnaeus). NMFS Sandy Hook Lab., Tech. Ser. Rept. No. 25. Robichaud-LeBlanc, K., S.C. Courtenay, and J.M. Hanson Ontogenetic diet shifts in age-0 striped bass, Morone saxatilis, from the Miramichi River estuary, Gulf of St. Lawrence. Can. J. Zool. 75: Roelofs, E.W Food studies of young sciaenid fishes, Micropogonias and Leiostomus, from North Carolina. Copeia 2:

68 Ross, S.T Resource partitioning in fish assemblages: a review of field studies. Copeia 1986(2): Rountree R.A Barracudas. Family Sphyraenidae. In: Collette, B.B., and G. Klein- MacPhee (eds.), Bigelow and Schroeder's Fishes of the Gulf of Maine, Smithsonian Institution, 3rd. Edition, Rountree, R.A. and K.W. Able Fauna of polyhaline subtidal marsh creeks in southern New Jersey: Composition, abundance and biomass. Estuaries 15(21): Rountree, R.A., and K.W. Able Seasonal abundance, growth, and foraging habits of juvenile smooth dogfish, Mustelus canis, in a New Jersey estuary. Fish. Bull., U.S. 94: Rudershausen, P Food, feeding and length-weight relationships of juvenile, young-of-the-year striped bass, Morone saxatilis and white perch, Morone americana, in the James River estuary, Virginia. M.S. thesis, College of William and Mary, School of Marine Science. 118 pp. Rulifson, R.A., and S.A. McKenna Food of striped bass in the upper Bay of Fundy, Canada. Trans. Amer. Fish. Soc.116: Ryer, C.H., and R.J. Orth Feeding ecology of the northern pipefish, Syngnathus fuscus, in a seagrass community of the lower Chesapeake Bay. Estuaries 10(4): Saloman, C.H., and S.P. Naughton Food of crevalle jack (Caranx hippos) from Florida, Louisiana, and Texas. NOAA Tech. Memo. NMFS-SEFC pp. Sand, R.L Aspects of the feeding ecology of the cunner, Tautogolabrus adspersus, in Narragansett Bay. M.S. thesis, Univ. of Rhode Island, Kingston. 94 pp. Sawyer, P.J Intertidal life-history of the rock gunnel, Pholis gunnellus, in the western Atlantic. Copeia 1967(1): Schaefer, R.H Feeding habits of striped bass from the surf waters of Long Island. N.Y. Fish Game J. 17(1):1-17. Schaeffer, J.S., and F.J. Margraf Food of white perch (Morone americana) and potential for competition with yellow perch (Perca flavescens) in Lake Erie. Ohio J. Sci. 86(1): Schmelz, G.W A natural history study of the mummichog, Fundulus heteroclitus (L.) in Canary Creek Marsh. M.S. thesis, University of Delaware, Newark, DE. 62

69 Schulze, M.B Using a field survey to assess potential temporal and spatial overlap between piscivores and their prey and a bioenergetics model to examine potential consumption of prey, especially juvenile anadromous fish, in the Connecticut River estuary. M.S. thesis, Univ. of Massachusetts. 133 pp. Schwartz, F.J. 1997b. Biology of the striped cusk-eel, Ophidion marginatum, from North Carolina. Bull. Mar. Sci. 61(2): Schwartz, F.J., and B.W. Dutcher Age, growth, and food of the oyster toadfish near Solomons, Maryland. Trans. Amer. Fish. Soc. 92(2): Scott W.B., and E.J. Crossman Freshwater fishes of Canada. Bull. Fish. Res. Bd. Can.184:966 pp. Scott, W.B., and M.G. Scott Atlantic fishes of Canada. Can. Bull. Fish. Aquat. Sci. 219:731 pp. Sedberry, G.R Food habits and trophic relationships of a community of fishes on the outer continental shelf. NOAA Tech. Rept. NMFS SSRF-773. Sedberry, G.R Food and feeding of black sea bass, Centropristis striata, in live bottom habitats in the south Atlantic Bight. J. Elisha Mitchell Sci. Soc. 104(2): Setzler, E.M., W.R. Boynton, K.V. Wood, H.H. Zion, L. Lubbers, N.K. Mountford, P. Frere, L. Tucker, and J.A. Mihursky Synopsis of biological data on striped bass, Morone saxatilis (Walbaum). NOAA Tech. Rept. NMFS Circ pp. Sheridan, P.F Food habits of the bay anchovy, Anchoa mitchilli, in Apalachicola Bay, Florida. Northeast Gulf Sci. 2(2): Sheridan, P.F Trophic resource utilization by three species of Sciaenid fishes in a northwest Florida estuary. Northeast Gulf Sci. 3(1):1-15. Sherman, K., and K.A. Honey Seasonal variation in the food of larval herring in coastal waters of central Maine. Cons. Perm. Explor. Mer 158: Sherman, K., and H.C. Perkins Seasonal variations in the food of juvenile herring in coastal waters of Maine. Trans. Amer. Fish. Soc. 100(1): Shipp, R.L., and R.W. Yerger Status, characters, and distribution of the northern and southern puffers of the genus Sphoeroides. Copeia 1969(3): Shumway, S.E., and R.R. Stickney Notes on the biology and food habits of the cunner. N.Y. Fish Game J. 22(1):

70 Siefert, R.E First food of larval yellow perch, white sucker, bluegill, emerald shiner, and rainbow smelt. Trans. Amer. Fish. Soc. 101(2): Sikora, W.B., R.W. Heard, and M.D. Dahlberg The occurrence and food habits of two species of hake, Urophycis regius and U. floridanus in Georgia estuaries. Trans. Amer. Fish. Soc Simmons, E.G., and J.P. Breuer A study of redfish, Sciaenops ocellata Linnaeus and black drum, Pogonias cromis Linnaeus. Publ. Inst. Mar. Sci. Univ. Tex. 8: Simpson, D.G., and G. Gunter Notes on the habitats, systematic characters, and life histories of Texas salt water Cyprinodontes. Tulane Stud. Zool. 4: Smith, L.D., and B.C. Coull Juvenile spot (Pisces) and grass shrimp poredation on meiobenthos in muddy and sandy substrata. J. Exp. Mar. Biol. Ecol. 105: Smith, H.M The fishes of North Carolina. N.C. Geol, and Economic Survey 2:xi, 449. Smith, D.G., and K. A. Tighe. 2002a. Freshwater eels. Family Anguillidae. In: Collette, B.B., and G. Klein-MacPhee (eds.), Bigelow and Schroeder's Fishes of the Gulf of Maine, Smithsonian Institution, 3rd. Edition pp. Smith, D.G., and K. A. Tighe. 2002b. Conger eels. Family Congridae. In: Collette, B.B., and G. Klein-MacPhee (eds.), Bigelow and Schroeder's Fishes of the Gulf of Maine, Smithsonian Institution, 3rd. Edition pp. Smith, K.J., G.L. Taghon, and K.W. Able Trophic linkages in marshes: ontogenetic changes in diet for young-of-the-year mummichog, Fundulus heteroclitus. In: M.P. Weinstein and D.A. Kreeger (eds.), Concepts and Controversies in Tidal Marsh Ecology. Kluwer Academic Publishers, The Netherlands pp. Smith, M.W Food of killifish and white perch in relation to supply. J. Fish. Res. Bd. Canada 7(1): Smith, R.W An analysis of the summer flounder, Paralichthys dentatus, population in the Delaware Bay. M.S. thesis, University of Delaware, Newark. Smith, R.W., and F.C. Daiber Biology of the summer flounder, Paralichthys dentatus, in Delaware Bay. Fish. Bull., U.S. 75: Smith, S.M., J.G. Hoff, S.P. O'Neil, and M.P. Weinstein Community and trophic organization of nekton utilizing shallow marsh habitats, York River, Virginia. Fish. Bull., U.S. 82(3):

71 Sorenson, P.W., M.L. Bianchini, and H.E. Winn Diel foraging activity of American eels, Anguilla rostrata (Lesueur), in a Rhode Island estuary. Fish. Bull., U.S. 84(3): Spraker, H., and H.M. Austin Diel feeding periodicity of Atlantic Silverside, Menidia menidia, in the York River, Chesapeake Bay, Virginia. J. Elisha Mitchell Sci. Soc. 113: Springer, V.G., and K.D. Woodburn An ecological study of the fishes of the Tampa Bay area. Fla. Board Conserv. Mar. Res. Lab. Prof. Pap. Ser. No pp. Stehlik, L.L., and C.J. Meise Diet of winter flounder in a New Jersey estuary: Ontogenetic change and spatial variation. Estuaries 23(3): Steimle, F.W., Jr., and L. Ogren Food of fish collected on artificial reefs in the New York Bight and off Charleston, South Carolina. Mar. Fish. Rev. 44(6-7): Steimle, F.W., D. Jeffress, S.A. Fromm, R.N. Reid, J.J. Vitaliano, and A. Frame Predator-prey relationships of winter flounder, Pleuronectes americanus, in the New York Bight apex. Fish. Bull., U.S. 92: Stevenson, R.A., Jr The biology of the anchovies Anchoa mitchilli mitchilli Cuvier and Valenciennes 1848 and Anchoa hepsetus hepsetus Linnaeus 1758 in Delaware Bay. M.A. thesis, University of Delaware, Newark, DE. Stickney, R.R., and R.B. McGeachin Food habits of fishes associated with marshland developed on dredged material. Proc. Ann. Conf. S.E. Assoc. Fish and Wildl. Agencies 32: Stickney, R.R., G.L. Taylor, and D.B. White Food habits of five species of young southeastern United States estuarine Sciaenidae. Chesapeake Sci. 16(2): Stone, H.H Composition, morphometric characteristics and feeding ecology of alewives (Alosa pseudoharengus) and blueback herring (Alosa aestivalis) (Pisces: Clupeidae) in Minas Basin. M.S. thesis, Acadia University, wolfville, Nova Scotia. 191pp. Stone, H.H. and G.R. Daborn Diet of Alewives, Alosa pseudoharengus and blueback herring, A. aestivalis (Pisces: Clupeidae) in Minas Basin, Nova Scotia, a turbid, macrotidal estuary. Envir. Biol. Fish. 19(1): Stone, H.H., and B.M. Jessop Feeding habits of anadromous alewives, Alosa pseudoharengus, off the Atlantic Coast of Nova Scotia. Fish. Bull., U.S. 92: Stoner, A.W., A.J. Bejda, J.P. Manderson, B.A. Phelan, L.L. Stehlik, and J.P. Pessutti. 65

72 1999. Behavior of winter flounder. Pseudopleuronectes americanus, during the reproductive season: laboratory and field observations on spawning, feeding, and locomotion. Fish. Bull., U.S. 97: Stouder, D.J., K.L. Fresh, and R.J. Feller (eds.) Theory and Application in Fish Feeding Ecology. University of South Carolina, Columbia, SC. 390 pp. Stroud, R.H Survey of the food of four fishes of the Bay of Fundy. Bowdoin Sci. Sta., Bowdoin College, 4th Ann. Rept. No. 6: Sumner, F.B., R.C. Osburn, and L.J. Cole A biological survey of the wates of Woods hole and vicinity. Part II., Sect. III. A catalogue of the marine fauna. Bull. U.S. Bur. Fish 31: Suzuki, K Biology of striped mullet Mugil cephalus Linnaeus. I. Food content of young. Rep. Fact. Prefect. Univ. Mie., 5: Taylor, E.T Food habits of dominant piscivorous fishes in Delaware Bay, with special reference to predation on juvenile weakfish. M.S. thesis, Univ. of Delaware Teixeira, R.L Reproductive and feeding biology of selected Syngnathids (Pisces: Teleostei) of the western Atlantic. Ph.D. dissertation, The College of William and Mary, Virginia. Thoits, C.F., III A compendium of the life history ecology of the white perch, Morone americana. Bull. Mass. Div. Fish and Game 24:1-16. Thomas, D.L The Early Life History and Ecology of Six Species of Drum (Sciaenidae) in the Lower Delaware River, a Brackish Tidal Estuary. Ichthyol. Assoc. Bull. 3 (PART III). 247 pp. Thompson, B.A Mullets. Family Mugilidae. In: Collette, B.B., and G. Klein- MacPhee (eds.), Bigelow and Schroeder's Fishes of the Gulf of Maine, Smithsonian Institution, 3rd. Edition, Timmons, M Relationships between macroalgae and juvenile fishes in the inland bays of Delaware. Ph.D. dissertation, Univ. of Delaware, Lewes, DE. Toepfer, C.S., and J.W. Fleeger Diet of juvenile fishes Citharichthys spilopterus, Symphurus plagiusa and Gobionellus boleosoma. Bull. Mar. Sci. 56(1): Trent, W.L., and W.W. Hassler Feeding behavior of adult striped bass, Roccus saxatilis, in relation to stages of sexual maturity. Chesapeake Sci. 7:

73 Tupper, M., and K.W. Able Movements and food habits of striped bass (Morone saxatilis) in Delaware Bay (USA) salt marshes: comparison of a restored and a reference marsh. Mar. Biol. 137: Urban, T.P., and S.B. Brandt Food and habitat partitioning between young-ofyear alewives and rainbow smelt in southeastern Lake Ontario. Envir. Biol. Fish. 36: Van Engle, W.A. and E.B. Joseph Characterization of coastal and estuarine fish nursery grounds as natural communities. U.S. Fish. Wildl. Serv. Commercial Fish. Res. and Devel. Act Final Rpt. Vazquez Rojas, A. V Energetics, trophic relationships and chemical composition of bay anchovy, Anchoa mitchilli in the Chesapeake Bay. M.S. thesis, University of Maryland, C.E.E.S., CBL, Solomons, MD. Vigerstad, T.J., and J.S. Cobb Effects of predation by sea-run juvenile alewives (Alosa pseudoharengus) on the zooplankton community at Hamilton Reservoir, Rhode Island. Estuaries 1(1): Vinogradov, V.I Studies of the food habits of silver and red hake in the northwest Atlantic area, ICNAF Research Bull. 9: Vivian, D.N., J.T. Duffy-Anderson, R.G. Arndt, and K.W. Able Feeding habits of young-of-the-year winter flounder, Pseudopleuronectes americanus, in the Hudson River estuary, U.S.A. Bull. N.J. Acad. Sci. 45(2):1-6. Vladykov, V.D Eels: Fishes of Canada. Quebec Dept. Fish. Album No pp. Walburg, C.H Observations of the food and growth of juvenile American shad Alosa sapidissima. Trans. Am. Fish. Soc 86: Walsh, G., and G.J. FitzGerald Resource utilization of three species of sticklebacks (Gasterostidae) in tidal salt-marsh pools. J. Fish Biol. 25: Walter, J.F., III Diet composition and feeding habits of large striped bass, Morone saxatilis in Chesapeake Bay. M.S. thesis, College of William and Mary, School of Marine Science. 123 pp. Walter, J.F., III, A.S. Overton, K.H. Ferry, and M.E. Mather Atlantic coast feeding habits of striped bass: a synthesis supporting a coast-wide understanding of trophic biology. Fish. Mgmt. Ecol. 10: Warkentine, B.E., and J.W. Rachlin Winter offshore diet of the Atlantic silverside, Menidia menidia. Copeia 1989(1):

74 Warlen, S.M Some aspects of the life history of Cyprinodon variegatus Lacepede 1803, in southern Delaware. Master s thesis, University of Delaware, Newark. Weaver, J.E Food selectivity, feeding chronology, and energy transformation of juvenile alewife (Alosa pseudoharengus) in the James River near Hopewell, Virginia. Ph.D. dissertation, University of Virginia, Charlottesville. Weinstein, M.P Shallowmarsh habitats as primary nurseries for fish and shellfish in the Cape Fear River, North Carolina. Fish. Bull., U.S. 77: Wells, B., D.H. Steele, and A.V. Tyler Intertidal feeding of winter flounders (Pseudopleuronectes americanus) in the Bay of Fundy. J. Fish. Res. Bd. Canada 30: Welsh, W.W., and C.M. Breder, Jr A contribution to the life history of the puffer, Sphoeroides maculatus (Schneider). Zoologica, N.Y. 2: Welsh, W.W., and C.M. Breder, Jr Contributions to the life histories of Sciaenidae of the eastern U.S. coast. Bull. U.S. Bur. Fish. 39: Wenner, C.A., and J.A. Musick Food habits and seasonal abundance of the American eel, Anguilla rostrata, from the Lower Chesapeake Bay. Chesapeake Sci. 16(1): Werme, C Resource partitioning in a saltmarsh fish community. Ph.D. dissertation, Boston University. 126 pp. Wilk, S.J Biological and fisheries data on bluefish, Pomatomus saltatrix (Linnaeus). NMFS Sandy Hook Lab., Tech. Ser. Rept. No. 11. Wilk, S.J Biological and fisheries data on weakfish, Cynoscion regalis (Bloch and Schnieder). NMFS Sandy Hook Lab., Tech. Ser. Rept., No pp. Williams, R.O., and G.E. Bruger Investigations on American Shad in the St. John's River. State of Florida Dept. Natural Resources Technical Series Report No pp. Wilson, C.A., J.M. Dean, and R. Radtke Age, growth rate and feeding habits of the oyster toadfish, Opsanus tau (Linnaeus) in South Carolina. J. Exp. Mar. Biol. Ecol. 62: Whoriskey, F.G Intertidal feeding and refuging by cunners, Tautogolabrus adspersus (Labridae). Fish. Bull., 81(2):

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76 LIST OF TABLES Table 1. Table 2a. Table 2b. Table 3. Table 3b. Table 4. Identification of prey ranking by young-of-the-year estuarine fish based on composite of studies examined Identification of prey ranking by young-of-the-year estuarine fish based on each study examined Details of specific studies which identified prey by young-of-the year estuarine fish Details of specific species which identified fish as prey by young-of- theyear estuarine fishes Identification of fish prey ranking by young-of-the-year estuarine fish based on each study examined. Size of young-of-the-year fish analyzed for stomach contents in the Middle Atlantic Bight based on the available literature LIST OF FIGURES Figure 1 Figure 2 Figure 3 Frequency distribution of number of food habit studies for individual fish species. Frequency distribution of number of fish stomach samples by fish species for each study. Species specific size distribution of young-of-the-year fishes examined during the studies included in this analysis. 70

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