Interannual Variation in Diet and Condition in Juvenile Bluefish During Estuarine Residency K

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Transactions of the American Fisheries Society Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/utaf20 Interannual Variation in Diet and Condition in Juvenile Bluefish during Estuarine Residency K. D. Friedland a , G. C. Garman b , A. J. Bejda c , A. L. Studholme c & B. Olla d a National Marine Fisheries Service, Northeast Fisheries Center, Woods Hole, Massachusetts, 02543, USA b Department of Biology, Virginia Commonwealth University, Richmond, Virginia, 23284, USA c National Marine Fisheries Service, Northeast Fisheries Center, Sandy Hook Laboratory, Highlands, New Jersey, 07732, USA d Cooperative Institute for Marine Resources Studies, National Marine Fisheries Service, Northwest Fisheries Center, Hatfield Marine Science Center, Newport, Oregon, 97365, USA Available online: 09 Jan 2011

To cite this article: K. D. Friedland, G. C. Garman, A. J. Bejda, A. L. Studholme & B. Olla (1988): Interannual Variation in Diet and Condition in Juvenile Bluefish during Estuarine Residency, Transactions of the American Fisheries Society, 117:5, 474-479 To link to this article: http:// dx.doi.org/10.1577/1548-8659(1988)117<0474:IVIDAC>2.3.CO;2

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Interannual Variation in Diet and Condition in Juvenile Bluefish during Estuarine Residency K. D. FRIEDLAND National Marine Fisheries Service, Northeast Fisheries Center Woods Hole. Massachusetts 02543,USA . GARMAC . G N Department of Biology, Virginia Commonwealth University Richmond. Virginia 23284, USA A. J. BEJDA AND A. L. STUDHOLME National Marine Fisheries Service, Northeast Fisheries Center Sandy Hook Laboratory. Highlands, JerseyNew 07732,USA . OLLB A Cooperative Institute Marinefor Resources Studies National Marine Fisheries Service. Northwest Fisheries Center Hatfield Marine Science Center. Newport, Oregon 97365, USA

Abstract.—We examined the diets and weight-length relationships of juvenile bluefish Poma- tomus saltatrix from Sandy Hook Bay, New Jersey, during 1981, 1983, and 1984. Diets consisted o varietfa polychaetef yo , crustacean fisd h,an prey. Opossum shrimp Neomysis americana. sand shrimp Crangon septemspinosa, grass shrimp Palaemonetes vulgar is, bay anchovy Anchoa mitchilli, striped killifish Fundulus majalis. Atlantid an c silverside Menidia menidia dominate diee dth t in term biomasf so frequencd san occurrencef yo . Consumptio invertebratf no fisd ehan prey varied between years. Bluefish condition factor was highest in 1981, when fish were the predominant prey lowed ,an 198n ri 1984d 3an , whe diete nth s consisted mostl invertebratf yo e prey diete .Th s of juvenile bluefish in Sandy Hook Bay contained more invertebrate prey than has been described previously.

Larval and early juvenile bluefish Pomatomus the basic information on patterns of annual vari- sallatrix distributee ar d alon continentae gth l shelf atio feedinn i growtd gan juvenilef ho addreso st s of the Atlantic coast of the USA from spring to the problem are lacking. The goal of our study was early summer (Kendall and Walford 1979). As to characterize juvenile bluefish diet and condi- they grow, they enter estuarie completo t s e tioth e n durin speciese gth oe f littoraus ' l zone nursery major portion of their first year's growth and de- habitats in Sandy Hook Bay, New Jersey, velopment (Clark 1973). Feeding and growth of juvenile bluefish, as with the estuarine young of the yea manf ro y species (Mille Dund ran n 1980), Methods Downloaded by [Oregon State University] at 16:32 02 September 2011 may increas estuarien ei s becaus hige e collecteth hf W epreo y d juvenile bluefish weekly from densities, high estuarine temperatures, and pro- Horseshoe Cove, a protected cove on the bay side tection from predators found there. Therefore, es- of Sandy Hook, New Jersey, from June to Septem- tuarine residency of the juveniles may be a critical her during 1981, 1983, and 1984. This area is in perio bluefisn di h life histor severar yfo l reasons polyhaline th , e portio Sandf no y Hoo (Draxy kBa - Year-class strength of bluefish, though probably ler et al. 1984). Fish were captured during daylight influenced most by environmental events at sea roughly 2 h after high tide with a 25-m beach seine (Norcros . 1974) al als y t affecte se e oe b ma , th (10-mm-ba y db r mesh burrpreserved % )an -10 a n i d variatio biotin n i abiotid can c conditions tha - in tere d solutio formaldehydf no seawatern ei . Within fluence the growth of young fish in estuaries. Poor 24 h of capture, we recorded the wet weight (0.01 growt causy hma e totadecreased an l) g lengtd - hsurvivaes (millimeters e th n i l eacf )o h fish, then tuary itsel r lea o fdecrease o dt d contributio o nt remove stomachl dal preserved san d the promn i - population fecundity by undersized fish. Neither panol for later analysis. Stomachs were opened phenomenon has been studied for bluefish, and individually, and the contents were identified to

474 JUVENILE BLUEFISH DIET 475

180 ^^

El50

CD 120

170 190 210 230 250 270 JULIAN DAY FIGURE 1.—Mean lengths (upper weightd an ) s (lower f juvenilo ) e bluefish collecte e Horseshoth t da e Cove sampling site, New Jersey. Error bars are ± 1 SD of the mean.

majoa r taxon, counted, , h) drie 4 t 60°2 d (a r Cfo uate between-year changes in relative condition. and weighed (to 0.001 g). Only length and weight data for fish with empty Downloaded by [Oregon State University] at 16:32 02 September 2011 We computed three food item indices to reduce stomachs were used for these comparisons be- the potential biases associated wit individuan ha l cause weight measurements were biase stomy db - measure (Hynes 1950; Windell 1971). The indices ach contents. Regressions of log^weight) against were (1) number of stomachs in which a taxon logXlength) were develope eacr dfo h year. Because occurred, expressed as a percentage of the total the slopethree th f es o regressions were homoge- numbe stomachf ro s containing percenfoo= F d( t neou 0.71)= P adjustee s( th , d mean f weighso t frequency of occurrence); (2) number of individ- were suitabl comparisor efo n with analysi- co f so ual eacf so h taxon, expresse percentaga s da f eo variance. This statistical test remove contrie dth - the total numbe foof ro d itempercen= N s- ( nu t bution of seasonal and ontogenetic effects upon merical abundance); and (3) weight of a taxon, the weight-length relationships (Le Cren 1951). expresse percentaga s da totae th lf eo weigh f o t food items (W = percent weight). Results We compared weight-length relationships - ofju diete juvenilf Th so e bluefis f varyino h g sizes venile bluefis r 1981hfo , 1983 d 198 an ,eval o 4t - (Figure 1) consisted of several invertebrate and 476 FRIEDLAN. AL T DE

TABLE 1.—Stomach contents of juvenile bluefish collected at Horseshoe Cove sampling site, New Jersey. F : percent frequenc occurrencef yo =;N percent numerical abundance percen= W ; t weight.

1981 1983 1984 Taxod nan sample size F N W F N W F N W Submillimcter prey Unidentified eggs 3.7 7.9 0.9 10.9 9.2 0.2 Unidentified copepods 3.9 69.4 0.4 Total 3.7 7.9 0.9 13.4 78.6 0.6 Polychaeia Nereis arenacedonta 1.2 0.0 0.7 Nereis virens 3.7 0.5 1.6 Nereis sp. 1.4 0.1 0.0 9.2 0.2 5.6 Glycera. sp 1.0 1.0 1.0 6.8 0.8 3.2 Unidentified Polychaeia 1.7 0.0 0.6 Total 1.0 1.0 1.0 11.5 1.4 4.8 11.9 0.2 6.9 Crustacea Neomysis americana 4.7 5.3 0.3 27.4 59.2 15.4 19.7 17.9 9.0 Crangon septemspinosa 26.4 32.0 8.6 42.2 12.1 23.2 26.1 0.8 19.6 Palaemonetes vu/garis 14.0 10.8 8.6 10.5 1.6 5.4 2.9 0.0 1.4 Gammarus spp. 2.1 10.6 0.6 2.4 0.4 0.1 5.8 0.2 0.3 Limulus polyphemus 0.7 0.1 0.1 0.3 0.3 0.4 Unidentified isopoda 0.2 0.0 0.0 Unidentified larvae 2.7 7.5 0.9 2.2 0.8 0.1 Unidentified Crustacea 3.1 3.8 0.6 0.7 0.0 0.7 6.5 0.1 0.4 Total 46.6 62.5 18.7 71.6 80.9 45.8 56.4 20.1 31.2 Telcostei Anchoa milchilli 9.8 4.0 17.7 19.6 2.5 40.9 17.8 0.2 42.6 h'undulus majalis 11.9 7.6 13.9 10.1 3.3 4.7 0.7 0.0 1.5 Fundulus Heteroclitus 1.0 0.0 1.5 0.7 0.0 0.3 0.3 0.0 2.8 Menidia menidia 34.2 18.9 38.3 1.7 0.2 0.7 4.1 0.0 4.8 Caranx hippos 0.3 0.0 0.0 Cynoscion regalis 2.1 1.8 5.4 Paralichthys dentatus 1.0 0.5 0.6 Unidentified larvae 2.7 2.5 0.8 0.3 0.0 0.1 Unidentified Teleostei 10.4 4.0 2.8 3.7 0.4 1.2 12.7 0.1 9.5 Total 66.3 36.8 80.2 36.8 8.9 48.6 36.3 0.3 61.3 Number of bluefish examined (% containing food) 193(84) (846 29 ) 589 (76)

fish prey. Prey that occurred in the bluefish diet valle jack Caranx hippos, weakfish Cynoscion re- 3 year l ial sn included polychaetes, opossum gait's, and summer flounder Paralichthys dentatus) shrimp Neomysis americana, sand shrimp Cran- occurred at low frequencies and were of little di- gon septemspinosa. grass shrimp Palaemonetes etary importance withi stude nth y area. vulgar is, gammarid amphipods, bay anchovy An- Whereas same manth f eyo pre y taxa occurred choa mitchilli, striped killifish Fundulus majalis, diee inth t each year feedine th , g indice thesr sfo e Downloaded by [Oregon State University] at 16:32 02 September 2011 mummichog Fundulus heteroclilus, and Atlantic taxa, especially in terms of the proportion of fish silversides Menidia menidia (Tabl thesf o ee e1)Us . invertebrato t e prey, varied among years 1981n .I , prey varied among yearssubsea t bu , t consisting fish dominated the diet; they occurred in 66% of of opossum shrimp, sand shrimp, grass shrimp, the stomachs r crustaceans, fo versu % 47 sd an , bay anchovy, striped killifish, and Atlantic silver- made up 80% of the diet by weight (Table 1). In diee th tf o % 79 sided an s , accounte90 , 87 r dfo contrast, during 198 d 19843an , fish were found by weight in 1981, 1983, and 1984, stomachse respectivelyth f o % ,36 respectively d . an 7 in3 , com- Unidentified eggs, copepods invertebratd ,an d ean pared with 72 and 56% for crustaceans, and made fish larvae were consumed onl Junn yi earld ean y diee weight61y d th b t %f an contri e o u 9 p4 Th . - July 198 d 19843an n 1984I . , high numberf so bution of polychaetes and submillimeter-size prey copepods were ingested, which lowere pere dth - to the 1981 diet was negligible; however, in 1983 cent-number index for other prey taxa. Through- and 1984 their contributions ranged from 4 to out the three sampling years, the remaining prey 13% by occurrence and from 1 to 7% by weight. items (horseshoe crab Limulus polyphemus, cre -between-yeae Th r patterns offis invertebratd han e JUVENILE BLUEFISH DIET 477

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60- 80- 100- 120- 140- 160- 70 90 110 130 150 170 BLUEFISH TOTAL LENGTH (mm) FIGURE 2.—Percent frequencie occurrencf so e (Ffisr )fo h (dotted lines invertebratd )an e prey (solid lines) eaten by juvenile bluefish, pooled by 10-mm predator length intervals. The number markers indicate study year (1 = 1981,3= 1983, an= 1984)d4 .

prey consumption were consistent over the size parisons of weights, adjusted for the covariate of rang f bluefiseo h studied. Percentage occurrence length, showed there were significant differences for fish pre y 10-myb m bluefish length intervals (P < 0.01) in the weight-length relationships be- was higher in 1981 than in either 1983 or 1984; tween years (Tabl Contrast. e2) adjustef so d means conversely e percenth , t occurrenc r invertefo e - showed the 1981 relationship to differ significant- brate prelowes y wa 198 n ri 1 tha 198n i r 1983o 4 ly from thos f 1980.035eo < P 3d 198( < )an P 4( (Figur . Further2) e , there wer o ontogenetin e c 0.001), whereas 198 198d 3an 4 regressiont no d sdi changes in consumption of fish versus inverte- differ significantly (P < 0.24) from each other. brate prey ove size rth e rang f bluefiseo ex e w h Weight s greate (IV)t lengta wa r 198) fo rh(L 1 amined. (log,** - '-12.16 3.127+ 8 log,/,) tha 198n ni 3 In addition to the different proportions of in- (log,Jf= -12.366 + 3.161 log,L)and 1984(log,tt' vertebrat fisd ehan diee preth t n amonyi g years, = -12.465 + 3.17810&X). there were also differences in the prey taxa most utilized mose Th . t important invertebrate taxf ao Discussion Downloaded by [Oregon State University] at 16:32 02 September 2011 the 1981 diet were sand shrim grasd pan s shrimp, diee juvenilf Th o t e bluefish from Sandy Hook based on frequency of occurrence and percent Bay contained a high proportion of invertebrates, weight (Figure 3). In contrast, in 1983 and 1984, which differs from results of previous bluefish diet sand shrimp and opossum shrimp were the two studies. Juvenile bluefish diet has been described most important invertebrate prey taxa grasd an , s qualitatively as consisting of a wide variety of prey, shrimlesf o ss importancepwa mose Th . t impor- including planktonic invertebrates, crustaceans, tant fish 198e preth 1n yi diet were descendinn i , g and fish (Bigelow and Schroeder 1953). Quanti- orde importancef ro , Atlantic silversides- an y ba , tative studies also indicate that juvenile bluefish chovy striped ,an d killifish (Figur contrastn I . e4) , consume a wide variety of prey, with fish domi- fish pre 198n y i d 1983an 4 diets were almos- ex t nant (Breder 1922; Grant 1962; Lassiter 1962; clusivel anchovyy yba , base frequencn do - oc f yo Smale and Kok 1983; Smale 1984). For example, currence and percent weight. Grant (1962) reported juvenile bluefish diets were Weight-length relationships for juvenile blue- 98% fish pre volumy yb e ove 2-yeara r studa n yi fish differed among the three study years. Com- Delaware estuary. In North Carolina, Lassiter 478 FRIEDLAND ET AL.

TABLE 2.—Analysi f covariancso f log-loeo g weight- length regressions for juvenile bluefish.

Sum of Mean Source df squares square F P Year 2 0.087 0.043 7.875 <0.01 Length 1 250.618 250.618 455.900.050 <0.01 Error 243 1.336 0.005 0.005

only 37 and 36% of the stomachs examined and contribute 61d %totaf an o 9 d4 l diet biomasy sb weight. Although difference specifin i s c prey taxa found in this study and elsewhere (for example, Atlantic sil versides in Sandy Hook Bay versus At- lantic menhaden Brevoortia tyrannus in the south- ern estuaries probable )ar site-specifi o t e ydu c difference availabilityn i s e higth , h proportiof o n benthic invertebrates in the diet suggests juvenile bluefish hav eflexibla e foraging behavior. Juvenile bluefish were heavie t lengta r h when FIGURE 3.—Percent frequencie occurrencf so d an ) e(F fish dominated their diets, suggestin grelationa - percent sanf weigho ) d shrim(W t p Crangon septemspi- ship between nursery diegrowthand t . Many fac- nosa, opossum shrimp Neomysis americana, grasd an s tors, such as temperature and competition, influ- shrimp Palaemonetes vulgaris in the diets of juvenile ence fish growth (Peters and Boyd 1972; Brett Jerseybluefisw Ne n .hi 1979; Bret Groved tan s wer1979)e w et unabl,bu e teso t t their effects withi contexe nth thif to s study. (1962) made nine monthly collections over 2 years , e 198Th 1 diet, dominate fishy db , must have con- and fish averaged 89% of the bluefish diets by tributed greatly to the observed differences in studyr year3 ou e f volumesth o , f fiso h2 pren I . y weigh t lengta t h amon years3 lowee e gth Th . r did not dominate bluefish diets, but occurred in weight-length relationship f 198 so 198d 3- an 4as sociated with predominantly invertebrate diets may be related to reduced caloric content of food items (Brett 1971; Thayer et al. 1973; Keast and Eadie 1985) or to higher energetic cost to the bluefish associated with foraging for and handling these prey (Mittlebach 1981; Soofiani and Hawkins 1982; Stein et al. 1984). effece Th t nursery growth rates could havn eo survival and fecundity at age makes the factors that control diet to be potentially important to the Downloaded by [Oregon State University] at 16:32 02 September 2011 recruitment process. More robust bluefish juveniles woul expectee db havo dt e lower ratef o s natural mortality during estuarine residency (Harden Jones 1981; Keas Eadid an t e 1985)o t , £ 26 better surviv migratioe eth n southwar fale th l n di (Ewing et al. 1984; Larsson 1985), and to be more productive spawners of subsequent year classes (DeAngelis and Coutant 1982). The combination of these factors could result in a higher contributio proximato nt subsequend ean t year classey sb FIGURE 4.—Percent frequencies of occurrence (F) and percent Atlantif weigho ) (W t c silversides Menidia me- juvenile bluefish from estuaries where fish dies i t nidia, anchovy ba y Anchoa mitchilli. striped an d killifish available. The conditions in estuaries that influ- Fundulus majalis in the diets of juvenile bluefish in New ence availabilit f fisyo h pre bluefiso yt h warrant Jersey. further investigation. JUVENILE BLUEFISH DIET 479

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