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Interannual Variation in Diet and
Condition in Juvenile Bluefish
during Estuarine Residency
a
b
c
K. D. Friedland , G. C. Garman , A. J. Bejda , A. L.
c
Studholme & 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
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Residency, Transactions of the American Fisheries Society, 117:5, 474-479
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Transactions of the American Fisheries Society 117:474-479, 1988
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
G. C. GARMAN
Department of Biology, Virginia Commonwealth University
Richmond. Virginia 23284, USA
Downloaded by [Oregon State University] at 16:32 02 September 2011
A. J. BEJDA AND A. L. STUDHOLME
National Marine Fisheries Service, Northeast Fisheries Center
Sandy Hook Laboratory. Highlands, New Jersey 07732, USA
B. OLLA
Cooperative Institute for Marine 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 Pomatomus saltatrix from Sandy Hook Bay, New Jersey, during 1981, 1983, and 1984. Diets consisted
of a variety of polychaete, crustacean, and fish prey. Opossum shrimp Neomysis americana. sand
shrimp Crangon septemspinosa, grass shrimp Palaemonetes vulgar is, bay anchovy Anchoa mitchilli, striped killifish Fundulus majalis. and Atlantic silverside Menidia menidia dominated the diet
in terms of biomass and frequency of occurrence. Consumption of invertebrate and fish prey varied
between years. Bluefish condition factor was highest in 1981, when fish were the predominant
prey, and lower in 1983 and 1984, when the diets consisted mostly of invertebrate prey. The diets
of juvenile bluefish in Sandy Hook Bay contained more invertebrate prey than has been described
previously.
Larval and early juvenile bluefish Pomatomus
sallatrix are distributed along the continental shelf
of the Atlantic coast of the USA from spring to
early summer (Kendall and Walford 1979). As
they grow, they enter estuaries to complete the
major portion of their first year's growth and development (Clark 1973). Feeding and growth of
juvenile bluefish, as with the estuarine young of
the year of many species (Miller and Dunn 1980),
may increase in estuaries because of the high prey
densities, high estuarine temperatures, and protection from predators found there. Therefore, estuarine residency of the juveniles may be a critical
period in bluefish life history for several reasons,
Year-class strength of bluefish, though probably
influenced most by environmental events at sea
(Norcross et al. 1974), may also be affected by the
variation in biotic and abiotic conditions that influence the growth of young fish in estuaries. Poor
growth may cause decreased survival in the estuary itself or lead to decreased contribution to
population fecundity by undersized fish. Neither
phenomenon has been studied for bluefish, and
the basic information on patterns of annual variation in feeding and growth of juveniles to address
the problem are lacking. The goal of our study was
to characterize juvenile bluefish diet and condition during the species' use of littoral zone nursery
habitats in Sandy Hook Bay, New Jersey,
Methods
We collected juvenile bluefish weekly from
Horseshoe Cove, a protected cove on the bay side
of Sandy Hook, New Jersey, from June to Septemher during 1981, 1983, and 1984. This area is in
the polyhaline portion of Sandy Hook Bay (Draxler et al. 1984). Fish were captured during daylight
roughly 2 h after high tide with a 25-m beach seine
(10-mm-bar mesh) and preserved in a 10% burrered solution of formaldehyde in seawater. Within
24 h of capture, we recorded the wet weight (0.01
g) and total length (millimeters) of each fish, then
removed all stomachs and preserved them in propanol for later analysis. Stomachs were opened
individually, and the contents were identified to
474
475
JUVENILE BLUEFISH DIET
180
^^
El50
CD 120
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90
45
170
190
210
230
250
270
JULIAN DAY
FIGURE 1.—Mean lengths (upper) and weights (lower) of juvenile bluefish collected at the Horseshoe Cove
sampling site, New Jersey. Error bars are ± 1 SD of the mean.
a major taxon, counted, dried (at 60°C for 24 h),
and weighed (to 0.001 g).
We computed three food item indices to reduce
the potential biases associated with an individual
measure (Hynes 1950; Windell 1971). The indices
were (1) number of stomachs in which a taxon
occurred, expressed as a percentage of the total
number of stomachs containing food (F = percent
frequency of occurrence); (2) number of individuals of each taxon, expressed as a percentage of
the total number of food items (N = percent numerical abundance); and (3) weight of a taxon,
expressed as a percentage of the total weight of
food items (W = percent weight).
We compared weight-length relationships of juvenile bluefish for 1981, 1983, and 1984 to eval-
uate between-year changes in relative condition.
Only length and weight data for fish with empty
stomachs were used for these comparisons because weight measurements were biased by stomach contents. Regressions of log^weight) against
logXlength) were developed for each year. Because
the slopes of the three regressions were homogeneous (P = 0.71), the adjusted means of weight
were suitable for comparison with analysis of covariance. This statistical test removed the contribution of seasonal and ontogenetic effects upon
the weight-length relationships (Le Cren 1951).
Results
The diets of juvenile bluefish of varying sizes
(Figure 1) consisted of several invertebrate and
476
FRIEDLAND ET AL.
TABLE 1.—Stomach contents of juvenile bluefish collected at Horseshoe Cove sampling site, New Jersey. F :
percent frequency of occurrence; N = percent numerical abundance; W = percent weight.
Taxon and
sample size
1981
F
N
1984
1983
W
F
N
W
F
N
W
3.7
7.9
0.9
3.7
7.9
0.9
10.9
3.9
13.4
9.2
69.4
78.6
0.2
0.4
0.6
1.2
0.0
0.7
0.5
0.1
0.8
1.6
0.0
3.2
9.2
0.2
5.6
1.7
11.9
0.0
0.2
0.6
6.9
19.7
26.1
2.9
5.8
0.3
0.2
2.2
6.5
56.4
17.9
0.8
0.0
0.2
0.3
0.0
0.8
0.1
20.1
9.0
19.6
1.4
0.3
0.4
0.0
0.1
0.4
31.2
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Submillimcter prey
Unidentified eggs
Unidentified copepods
Total
Polychaeia
Nereis arenacedonta
Nereis virens
Nereis sp.
Glycera sp.
Unidentified Polychaeia
Total
Crustacea
Neomysis americana
Crangon septemspinosa
Palaemonetes vu/garis
Gammarus spp.
Limulus polyphemus
Unidentified isopoda
Unidentified larvae
Unidentified Crustacea
Total
Telcostei
Anchoa milchilli
h'undulus majalis
Fundulus Heteroclitus
Menidia menidia
Caranx hippos
Cynoscion regalis
Paralichthys dentatus
Unidentified larvae
Unidentified Teleostei
Total
Number of bluefish examined
(% containing food)
1.0
1.0
1.0
3.7
1.4
6.8
1.0
1.0
1.0
11.5
1.4
4.8
4.7
26.4
14.0
2.1
5.3
32.0
10.8
10.6
0.3
8.6
8.6
0.6
27.4
42.2
10.5
2.4
0.7
59.2
12.1
1.6
0.4
0.1
15.4
23.2
5.4
0.1
0.1
3.1
46.6
3.8
62.5
0.6
18.7
2.7
0.7
71.6
7.5
0.0
80.9
0.9
0.7
45.8
9.8
11.9
1.0
34.2
4.0
7.6
0.0
18.9
17.7
13.9
1.5
38.3
19.6
10.1
0.7
1.7
2.5
3.3
0.0
0.2
40.9
4.7
0.3
0.7
17.8
0.7
0.3
4.1
0.3
0.2
0.0
0.0
0.0
0.0
42.6
1.5
2.8
4.8
0.0
2.1
1.0
1.8
0.5
5.4
0.6
10.4
66.3
4.0
36.8
2.8
80.2
2.7
3.7
36.8
2.5
0.4
8.9
0.8
1.2
48.6
0.3
12.7
36.3
0.0
0.1
0.3
0.1
9.5
61.3
193(84)
fish prey. Prey that occurred in the bluefish diet
in all 3 years included polychaetes, opossum
shrimp Neomysis americana, sand shrimp Crangon septemspinosa. grass shrimp Palaemonetes
vulgar is, gammarid amphipods, bay anchovy Anchoa mitchilli, striped killifish Fundulus majalis,
mummichog Fundulus heteroclilus, and Atlantic
silversides Menidia menidia (Table 1). Use of these
prey varied among years, but a subset consisting
of opossum shrimp, sand shrimp, grass shrimp,
bay anchovy, striped killifish, and Atlantic silversides accounted for 87, 90, and 79% of the diet
by weight in 1981, 1983, and 1984, respectively.
Unidentified eggs, copepods, and invertebrate and
fish larvae were consumed only in June and early
July 1983 and 1984. In 1984, high numbers of
copepods were ingested, which lowered the percent-number index for other prey taxa. Throughout the three sampling years, the remaining prey
items (horseshoe crab Limulus polyphemus, cre-
296 (84)
589 (76)
valle jack Caranx hippos, weakfish Cynoscion regait's, and summer flounder Paralichthys dentatus)
occurred at low frequencies and were of little dietary importance within the study area.
Whereas many of the same prey taxa occurred
in the diet each year, the feeding indices for these
taxa, especially in terms of the proportion of fish
to invertebrate prey, varied among years. In 1981,
fish dominated the diet; they occurred in 66% of
the stomachs, versus 47% for crustaceans, and
made up 80% of the diet by weight (Table 1). In
contrast, during 1983 and 1984, fish were found
in 37 and 36% of the stomachs, respectively, compared with 72 and 56% for crustaceans, and made
up 49 and 61% of the diet by weight. The contribution of polychaetes and submillimeter-size prey
to the 1981 diet was negligible; however, in 1983
and 1984 their contributions ranged from 4 to
13% by occurrence and from 1 to 7% by weight.
The between-year patterns offish and invertebrate
477
JUVENILE BLUEFISH DIET
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U_K)0
6070
8090
100110
120130
140150
160170
BLUEFISH TOTAL LENGTH (mm)
FIGURE 2.—Percent frequencies of occurrence (F) for fish (dotted lines) and invertebrate prey (solid lines) eaten
by juvenile bluefish, pooled by 10-mm predator length intervals. The number markers indicate study year (1 =
1981,3= 1983, and 4= 1984).
prey consumption were consistent over the size parisons of weights, adjusted for the covariate of
range of bluefish studied. Percentage occurrence length, showed there were significant differences
for fish prey by 10-mm bluefish length intervals (P < 0.01) in the weight-length relationships bewas higher in 1981 than in either 1983 or 1984; tween years (Table 2). Contrasts of adjusted means
conversely, the percent occurrence for inverte- showed the 1981 relationship to differ significantbrate prey was lower in 1981 than in 1983 or 1984 ly from those of 1983 (P < 0.035) and 1984 (P <
(Figure 2). Further, there were no ontogenetic 0.001), whereas 1983 and 1984 regressions did not
changes in consumption of fish versus inverte- differ significantly (P < 0.24) from each other.
brate prey over the size range of bluefish we ex- Weight (IV) at length (L) was greater for 1981
amined.
(log,**' - -12.167 + 3.128 log,/,) than in 1983
In addition to the different proportions of in- (log,Jf= -12.366 + 3.161 log,L)and 1984(log,tt'
vertebrate and fish prey in the diet among years, = -12.465 + 3.17810&X).
there were also differences in the prey taxa most
Discussion
utilized. The most important invertebrate taxa of
the 1981 diet were sand shrimp and grass shrimp,
The diet of juvenile 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, and grass qualitatively as consisting of a wide variety of prey,
shrimp was of less importance. The most impor- including planktonic invertebrates, crustaceans,
tant fish prey in the 1981 diet were, in descending and fish (Bigelow and Schroeder 1953). Quantiorder of importance, Atlantic silversides, bay an- tative studies also indicate that juvenile bluefish
chovy, and striped killifish (Figure 4). In contrast, consume a wide variety of prey, with fish domifish prey in 1983 and 1984 diets were almost ex- nant (Breder 1922; Grant 1962; Lassiter 1962;
clusively bay anchovy, based on frequency of oc- 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 prey by volume over a 2-year study in a
fish differed among the three study years. Com- Delaware estuary. In North Carolina, Lassiter
478
FRIEDLAND ET AL.
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TABLE 2.—Analysis of covariance of log-log weightlength regressions for juvenile bluefish.
FIGURE 3.—Percent frequencies of occurrence (F) and
percent weight (W) of sand shrimp Crangon septemspinosa, opossum shrimp Neomysis americana, and grass
shrimp Palaemonetes vulgaris in the diets of juvenile
bluefish in New Jersey.
(1962) made nine monthly collections over 2 years,
and fish averaged 89% of the bluefish diets by
volume. In 2 of the 3 years of our study, fish prey
did not dominate bluefish diets, but occurred in
£ 26
FIGURE 4.—Percent frequencies of occurrence (F) and
percent weight (W) of Atlantic silversides Menidia menidia, bay anchovy Anchoa mitchilli. and striped killifish
Fundulus majalis in the diets of juvenile bluefish in New
Jersey.
Source
df
Sum of
squares
Mean
square
F
P
Year
Length
Error
2
1
243
0.087
250.618
1.336
0.043
250.618
0.005
7.875
455.900.050
0.005
<0.01
<0.01
only 37 and 36% of the stomachs examined and
contributed 49 and 61% of total diet biomass by
weight. Although differences in specific prey taxa
found in this study and elsewhere (for example,
Atlantic sil versides in Sandy Hook Bay versus Atlantic menhaden Brevoortia tyrannus in the southern estuaries) are probably due to site-specific differences in availability, the high proportion of
benthic invertebrates in the diet suggests juvenile
bluefish have a flexible foraging behavior.
Juvenile bluefish were heavier at length when
fish dominated their diets, suggesting a relationship between nursery diet and growth. Many factors, such as temperature and competition, influence fish growth (Peters and Boyd 1972; Brett
1979; Brett and Groves 1979), but we were unable
to test their effects within the context of this study.
The 1981 diet, dominated by fish, must have contributed greatly to the observed differences in
weight at length among the 3 years. The lower
weight-length relationships of 1983 and 1984 associated 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).
The effect nursery growth rates could have on
survival and fecundity at age makes the factors
that control diet to be potentially important to the
recruitment process. More robust bluefish juveniles would be expected to have lower rates of
natural mortality during estuarine residency
(Harden Jones 1981; Keast and Eadie 1985), to
better survive the migration southward in the fall
(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 contribution to proximate and subsequent year classes by
juvenile bluefish from estuaries where fish diet is
available. The conditions in estuaries that influence availability of fish prey to bluefish warrant
further investigation.
JUVENILE BLUEFISH DIET
Acknowledgments
We lhank D. Kent, S. Maurawsky, V. Krouse,
G. Loftlin, C. Samet, and M. Cox for their help
in completing this study.
Downloaded by [Oregon State University] at 16:32 02 September 2011
References
Bigelow, H. B., and W. C. Schroeder. 1953. Fishes of
the Gulf of Maine. U.S. Fish and Wildlife Service
Fishery Bulletin 53(74).
Breder, C. M. 1922. Observations on young bluefish.
Copeia 1922:34-36.
Brett, J. R. 1971. Growth responses of young sockeye
salmon (Oncorhynchus nerka) to different diets and
planes of nutrition. Journal of the Fisheries Research Board of Canada 28:1635-1643.
Brett, J. R. 1979. Environmental factors and growth.
Pages 599-625 in W. S. Hoar, D. J. Randall, and
J. R. Brett, editors. Fish physiology, volume 8. Academic Press, New York.
Brett, J.R., and T. D. D. Groves. 1979. Physiological
energetics. Pages 280-344 in W. S. Hoar, D. J. Randall, and J. R. Brett, editors. Fish physiology, volume 8. Academic Press, New York.
Clark, J. R. 1973. Bluefish. Pages 250-251 in A. L.
Pacheco, editor. Proceedings of a workshop on egg,
larval and juvenile stages of fish in Atlantic coast
estuaries. National Marine Fisheries Service, Middle Atlantic Coast Fisheries Center, Technical Publication 1, Highlands, New Jersey.
DeAngelis, D. L., and C. C. Coutant. 1982. Genesis of
bimodal size distributions in species cohorts. Transactions of the American Fisheries Society 111:384388.
Draxler, A. F. J., R. Waldhauer, A. Matte, and J. B.
Mahoney. 1984. Nutrients, hydrography and their
relationship to phytoflagellates in the Hudson-Raritan estuary. Bulletin New Jersey Academy of Science 29:97-179.
Ewing, R. D., C. E. Hart, C. A. Fustish, and G. Concannon. 1984. Effects of size and time of release
on seaward migration of spring chinook salmon,
Oncorhynchus tshawytscha. U.S. National Marine
Fisheries Service Fishery Bulletin 82:157-164.
Grant, G. C. 1962. Predation of bluefish on young
Atlantic menhaden in Indian River, Delaware.
Chesapeake Science 3:45-46.
Harden Jones, F. R. 1981. Fish migration: strategy and
tactics. Pages 139-165 in D. J. Aidley, editor. Animal migration. Cambridge University Press, New
York.
Hynes, H. B. N. 1950. The food of fresh-water sticklebacks (Gasterosteus aculatus and Pygosteus pungitius), with a review of methods used in studies of
the food of fishes. Journal of Animal Ecology 19:
36-58.
Keast, A., and J. M. Eadie. 1985. Growth dispensation
in year-0 largemouth bass, the influence of diet.
Transactions of the American Fisheries Society 114:
204-213.
479
Kendall,A.W.,andL.W.Walford. 1979. Sources and
distribution of bluefish, Pomatomus saltatrix, larvae and juveniles off the east coast of the United
States. U.S. National Marine Fisheries Service Fishery Bulletin 77:213-227.
Larsson, P. O. 1985. Predation on migrating smolt as
a regulating factor in Baltic salmon, Sal mo salar L.,
populations. Journal of Fish Biology 26:391-397.
Lassiter, R. R. 1962. Life history aspects of the bluefish, Pomatomus saltatrix (Linnaeus), from the coast
of North Carolina. Master's thesis. North Carolina
State University, Raleigh.
Le Cren, E. D. 1951. The length-weight relationships
and seasonal cycle in gonad weight and condition
in the perch (Perca fluviatilis). Journal of Animal
Ecology 20:201-219.
Miller, J.M., and M. L.Dunn. 1980. Feeding strategies
and patterns of movements in juvenile estuarine
fishes. Pages 437-448 in V. S. Kennedy, editor. Estuarine perspectives. Academic Press, New York.
Mittlebach, G. G. 1981. Foraging efficiency and body
size: a study of optimal diet and habitat use by bluegills. Ecology 62:1370-1386.
Norcross, J. J., S. L. Richardson, W. H. Massman, and
E. B. Joseph. 1974. Development of young bluefish (Pomatomus saltatrix) and distribution of eggs
and young in Virginian coastal waters. Transactions
of the American Fisheries Society 103:477-497.
Peters, D. S., and M. T. Boyd. 1972. The effect of
temperature, salinity and availability of food on the
feeding and growth of the hogchoker, Trinectes
maculatus (Block and Schneider). Journal of Experimental Marine Biology and Ecology 9:201-207.
Smale, M.J. 1984. Inshore small-mesh trawling survey
of the Cape south coast. Pan 3. The occurrence and
feeding of Argyrosomus hololepidotus, Pomatomus
saltatrix and Merluccius capensis. South African
Journal of Zoology 19:170-179.
Smale, M. J., and H. M. Kok. 1983. The occurrence
and feeding of Pomatomus saltatrix (elf) and Lichia
amia (leervis) juveniles in two Cape south estuaries.
South African Journal of Zoology 18:337-342.
Soofiani, N. M., and A. D. Hawkins. 1982. Energetic
costs at different levels of feeding in juvenile cod,
Gadus morhua L. Journal of Fish Biology 21:577592.
Stein, R. A., C. G. Goodman, and E. A. Marschall. 1984.
Using time and energetic measures of cost in estimating prey value for fish predators. Ecology 65:
702-715.
Thayer, G. W., W. E. Schaaf, J W. Angelovic, and M.
W. LaCroix. 1973. Caloric measurement of some
estuarine organisms. U.S. National Marine Fisheries Service Fishery Bulletin 71:289-296.
Windell, J. T. 1971. Food analysis and rate digestion.
IBP (International Biological Programme) Handbook 3:215-226.
Received December 3, 1987
Accepted November I, 1988