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03 Buckel FISH 97(4) 758 Abstract.–After spending summer Foraging habits of bluefish, months in estuaries, spring- and sum- mer-spawned young-of-the-year (YOY) Pomatomus saltatrix, on the bluefish, Pomatomus saltatrix, migrate out to continental shelf waters of the U.S. east coast continental shelf* Mid-Atlantic Bight in early autumn. Adult bluefish are found on the conti- nental shelf throughout summer and Jeffrey A. Buckel fall. Both juveniles and adults have high Marine Sciences Research Center food consumption rates and are gener- State University of New York ally piscivorous. To determine princi- Stony Brook, New York 11794-5000 pal prey types on the shelf, dietary Present address: James J. Howard Marine Sciences Laboratory analyses were performed on YOY and National Marine Fisheries Service, NOAA adult bluefish collected from National 74 Magruder Road Marine Fisheries Service autumn bot- Highlands, New Jersey 07732 tom trawl surveys in 1994 and 1995. E-mail address: [email protected] Both spring- and summer-spawned YOY bluefish diets were dominated by bay anchovy. However, the significantly Michael J. Fogarty larger size of the spring-spawned cohort Northeast Fisheries Science Center was associated with the consumption National Marine Fisheries Service, NOAA of other prey species such as squid, 166 Water Street butterfish, striped anchovy, and round Woods Hole, Massachusetts 02543 herring. Summer-spawned bluefish were significantly smaller in 1995 than in 1994; diet and prey size comparisons David O. Conover suggest that body size had a dramatic Marine Sciences Research Center influence on the amount of piscivorous State University of New York feeding in the summer-spawned cohort. Stony Brook, New York 11794-5000 Adult bluefish diet was dominated by schooling species such as squid, butter- fish, and clupeids. Cannibalism was virtually nonexistent. Daily ration es- timates of YOY bluefish on the shelf (4– 12% body wt/d) were similar to estua- One of the dominant marine pisci- striped bass during their summer– rine estimates in late summer. It is es- vores along the U.S. east coast is the fall growing season (Buckel et al., timated that during the month of Sep- bluefish, Pomatomus saltatrix 1999). After spending the summer tember, YOY bluefish in aggregate con- (Juanes et al., 1996). Bluefish months in estuaries, YOY bluefish sumed 6.0 to 6.8 billion bay anchovies in 1994 and from 2.2 to 5.3 billion in spawned in offshore waters of the migrate back out onto the shelf and 1995. The effect of this predatory loss South Atlantic Bight in the spring migrate southward to overwinter on population dynamics of bay anchovy (spring-spawned) are advected north- (Munch and Conover, in press). and the fish community on the conti- ward in waters associated with the Spring-spawned YOY bluefish mi- nental shelf is unknown. Gulf Stream (Hare and Cowen, 1996) grate out of estuaries at a size of and move into mid-Atlantic Bight ~180 mm fork length (FL) and ~100 g estuaries in June at ~60 mm fork (Nyman and Conover, 1988; McBride length (Kendall and Walford, 1979; and Conover 1991). By age 1, these Nyman and Conover, 1988; McBride spring-spawned fish attain sizes of and Conover, 1991). A second wave ~260 mm FL and weights of ~300 g of recruits consisting of summer- (Chiarella and Conover, 1990). spawned fish occur in nearshore wa- Therefore, an individual spring- ters from mid to late summer. spawned bluefish gains from 100 to In estuaries, bluefish exhibit 200 g during its autumn migration. rapid growth rates that are fueled Given a 15% gross growth efficiency by high food consumption and (Juanes and Conover, 1994; Buckel evacuation rates (Juanes and Con- et al. 1995), a single bluefish could over, 1994; Buckel et al., 1995; Buckel thus consume from ~650 to 1300 g and Conover, 1996). In the Hudson of prey. River estuary, for example, preda- tion by bluefish is high enough to * Contribution 1134 of the Marine Sciences Manuscript accepted 1 December 1998. account for virtually all natural mor- Research Center, State University of New Fish. Bull. 97:758–775 (1999). tality of young-of-the-year (YOY) York, Stony Brook, New York 11794. Buckel et al.: Foraging habits of Pomatomus saltatrix 759 Compared with the nearshore phase of the early Munch and Conover’s (in press) geographical bound- life history of bluefish, our knowledge of the forag- aries for this analysis. In 1994, summer-spawned blue- ing ecology and predatory impact of bluefish on the fish sizes by region were as follows: SOC, FL≤120 mm; continental shelf is poor. Here we quantify the diet C–D, FL≤160 mm; and SNE, FL≤160 mm. In 1995, of YOY and adult bluefish and determine YOY blue- summer-spawned bluefish sizes were as follows: fish prey type and size selectivity patterns, foraging SOC, FL≤100 mm; C–D, FL≤150 mm; and SNE, chronology, daily ration, and biomass of prey con- FL≤140 mm. Spring-spawned bluefish were those sumed during the autumn migration on the shelf. fish larger than the summer-spawned cohort by re- gion but ≤300 mm FL. Diets of spring- and summer-spawned juveniles Methods and adult bluefish were quantified. Bluefish taken for stomach content analysis were wet weighed (±1.0 g) Study area and collections and measured for fork length, FL (to 1.0 mm). Stom- achs were removed at sea and preserved in 10% for- YOY and adult bluefish and their potential prey were malin buffered with sea water. On some occasions, collected on the U.S. east coast continental shelf dur- whole fish were either frozen or preserved in 10% for- ing the autumn of 1994 and 1995 aboard the research malin and then processed in the laboratory. Stomach vessel Albatross IV. Collections were made during contents of bluefish were identified to the lowest pos- the National Marine Fisheries Service, Northeast sible taxon, enumerated, blotted dry, weighed (±0.01 g), Fisheries Science Center’s (NMFS-NEFSC) autumn and measured (TL for fish prey and mantle length for bottom trawl survey cruises at predetermined sta- squid, ±1.0 mm). Eye diameter (±0.1 mm) and caudal tions from Cape Hatteras, NC, to Nova Scotia. peduncle height (±0.1 mm) were measured for partially Cruises began in early September and ended in late eaten bay anchovy and butterfish prey and converted October in both years (6 September to 28 October into TL from linear regression equations (Scharf et al., 1994 and 5 September to 27 October 1995). 1997; Scharf et al., 1998a). Descriptions of the survey design and the trawl Each trawl containing bluefish provided us with a characteristics can be found in Azarovitz (1981). group or “cluster” of bluefish for a given station. Briefly, tows were made with a no. 36 Yankee trawl Therefore, for our analysis, the mean and variance equipped with rollers with an opening 3.2 m high, of diet indices were calculated with cluster sampling 10.4 m wide, with 12.7-cm stretched mesh in the estimators developed at the NEFSC-NMFS, Woods opening, with 11.4-cm stretched mesh in the codend, Hole, MA (Cochran, 1977, Fogarty, unpubl. data). and with a 1.25-cm stretched mesh lining in the These calculations are described in a previous study codend and upper belly to retain YOY fishes. Tows that examined bluefish diet in the Hudson River es- were 30 minutes in duration at 3.5 knots in relation tuary (Buckel et al., 1999). to bottom and were conducted on a 24-h basis. Net feeding Dietary analyses During a 30-minute tow, bluefish may feed within Adult and YOY bluefish were distinguished by us- the trawl (an activity known as net feeding). Net feed- ing length-age relationships. Munch and Conover (in ing could bias the diet index estimates or affect gut press) examined the annual size distributions of blue- fullness level estimates (or do both). We tested for fish from autumn bottom trawl surveys conducted such bias by classifying fish or squid prey as either since the 1970s in four different regions of the shelf: 1) “fresh” or 2) “digested” during our stomach con- SOC=South of Chesapeake Bay; C–D=Chesapeake tent examination. “Fresh” prey had no sign of diges- Bay to Delaware Bay; SNE=Southern New England tion and “digested” prey were either partially or well (Delaware Bay to Narragansett Bay); and Georges digested (e.g. they were anywhere from starting to Bank. On the basis of these size distributions and lose skin to being only identifiable by skeleton or backcalculated sizes at age 1 (Chiarella and Conover, shape). The percent occurrence of four dominant prey 1990, and references therein), they classified YOY (bay anchovy, striped anchovy, butterfish, and squid) bluefish in autumn as those fish ≤300 mm FL. We were compared between fresh and digested catego- classified adults as those bluefish >300 mm FL and ries in YOY bluefish. If the percent occurrence of each we distinguished spring- and summer-spawned YOY prey category within a cohort and region was simi- bluefish for each region in 1994 and 1995 on the ba- lar for fresh and digested prey, it would suggest that sis of bimodality in length-frequency distributions either there was no net feeding or that it did not af- (Munch and Conover, in press). We also adopted fect diet index estimates. 760 Fishery Bulletin 97(4), 1999 Prey-type selectivity Size-selective feeding on bay anchovy prey was examined by comparing the sizes of bay anchovy in- The feeding selectivity of spring-spawned bluefish gested by bluefish at a specific station with sizes of on the continental shelf was determined from the bay anchovy captured in the trawl at that station.
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