Lane et al.: Ontogenetic and temporal variability in the fat content and fatty acid composition of Clupea harengus 
121 
on which size of fish is eaten, and which season and 
year it is consumed in. The current study demonstrates 
the variability in the nutritional quality of species that 
may be assumed to offer consistent value in food webs 
and underscores the importance of characterizing spe- 
cies over broad temporal and developmental scales to 
capture this variability. 
Acknowledgments 
The authors thank the following, without whom this 
project would not have been possible: the fishermen of 
Grand Manan for herring samples, especially the Foster 
and Kinghorne families; J. Fife, G. Melvin, R. Stephen- 
son, M. Powers (all from the Canadian Department of 
Fisheries and Ocean, St. Andrews Biological Station), 
L. Murison, and the Grand Manan Whale and Seabird 
Research Station staff for logistical and intellectual 
support; T. Hooper, D. Cooke, and K. Dugan at Connors 
Brothers, Inc., for sample collection; D. P. Gannon and 
F. Scharf for project development and assistance; and 
S. Budge for statistical support and guidance and for 
confirmation of peak identification. This work was sup- 
ported through a Department of Fisheries and Oceans 
Herring Science Council research grant to H.A.L, and 
by the Grand Manan Whale and Seabird Research Sta- 
tion and the University of North Carolina Wilmington. 
Literature cited 
Alonso-Alvarez, C., and J. L. Telia. 
2001. Effects of experimental food restriction and body 
mass changes on the avian T-cell-mediated immune 
response. Can. J. Zool. 79:101-105. 
Anthony, V. C. 
1972. Population dynamics of the Atlantic herring in the 
Gulf of Maine. Ph.D. diss., 266 p. Univ. Washington, 
Seattle, WA. 
Anthony, V. C., and G. Waring. 
1980. The assessment and management of the Georges 
Bank herring fishery. Rapp. P.-V. Reun.-Cons. Int. 
Explor. Mer 177:72-111. 
Atkinson, S. N., and M. A. Ramsay. 
1995. The effects of prolonged fasting of the body com- 
position and reproductive success of female polar bears 
( TJrsus maritimus). Funct. Ecol. 9:559-567. 
Bishop, D. G., D. A. Ritz, G. W. Hosie, J. R. Kenrick and J. Olley. 
1983. Fatty acid composition of the lipids of Puffinus 
tenuirostris (Temminck) in relation to its diet. J. Exp. 
Mar. Biol. Ecol. 71:17-26. 
Boyar, H. C., R. R. Marak, F. E. Perkins, and R. A. Clifford 
1973. Seasonal distribution and growth of larval her- 
ring ( Clupea harengus L.) in the Georges Bank-Gulf 
of Maine area from 1962-1970. J. Cons. Int. Explor. 
Mer 35:36-51. 
Bradford, R. G. 
1993. Differential utilization of storage lipids and stor- 
age proteins by Northwest Atlantic herring ( Clupea 
harengus harengus). J. Fish. Biol. 43:811-824. 
Budge, S. M., S. J. Iverson, W. D. Bowen, and R. G. Ackman. 
2002. Among- and within-species variability in fatty 
acid signatures of marine fish and invertebrates on 
the Scotian Shelf, Georges Bank, and southern Gulf of 
St. Lawrence. Can. J. Fish. Aquat. Sci. 59:886-898. 
Budge, S. M., S. J. Iverson, and H. N. Koopman. 
2006. Studying trophic ecology in marine ecosystems 
using fatty acids: A primer on analysis and interpreta- 
tion. Mar. Mamm. Sci. 22:759-801. 
Burnham, K. P., and D. R. Anderson. 
2002. Model selection and inference: a practical 
approach. Springer, New York. 
Chase, B. C. 
2002. Differences in diet of Atlantic tuna (Thunnus 
thynnus) at five seasonal feeding grounds of the New 
England continental shelf. Fish. Bull. 100:168-180. 
Clarke, K. R. 
1993. Non-parametric multivariate analyses of changes 
in community structure. Aust. J. Ecol. 18:117-143. 
Clarke, K. R., and R. N. Gorley. 
2006. PRIMER, vers. 6: user manual/tutorial, 190 p. 
PRIMER-E: Plymouth, UK. 
Clarke, K. R., and R. M. Warwick. 
2001. Changes in marine communities: an approach 
to statistical analysis and interpretation, 2 nd ed., 172 
p. PRIMER-E: Plymouth, UK. 
Colwell, M. A., and S. L. Landrum. 
1993. Nonrandom shorebird distribution and fine-scale 
variation in prey abundance. Condor. 95:94—103. 
Colton, J. B., Jr., W. G. Smith, A. W. Kendall Jr., P. L. Berrien, 
and M. P. Fahay. 
1979. Principle spawning areas and times of marine 
fishes, Cape Sable to Cape Hatteras. Fish. Bull. 
76:911-915. 
Dalsgaard, J., M. St. John, G. Kattner, D. Muller-Navarra, and 
W. Hagen. 
2003. Fatty acid trophic markers in the pelagic marine 
environment. Adv. Mar. Biol. 46:225-340. 
De Silva, S. S. 
1973. Food and feeding habits of the herring Clupea 
harengus and the sprat C. sprattus in inshore waters 
of the west coast of Scotland. Mar. Biol. 20:282-290. 
Diamond, A. W., and C. M. Devlin. 
2003. Seabirds as indicators of changes in marine 
ecosystems: ecological monitoring on Machias Seal 
Island. Environ. Monit. Assess. 88:153-175. 
Duffy, J. E., and V. J. Paul. 
1992. Prey nutritional quality and the effectiveness of 
chemical defenses against tropical reef fishes. Oeco- 
logia 90:333-339. 
Folch, J., M. Lees, and G. H. Sloane-Stanley. 
1957. A simple method for the isolation and purifica- 
tion of total lipids from animal tissues. J. Biol. Chem. 
226:497-509. 
Forsman, A., and L. E. Lindell. 
1997. Responses of a predator to variation in prey abun- 
dance: survival and emigration of adders in relation to 
vole density. Can. J. Zool. 75:1099-1108. 
Gannon, D. P., J. E. Craddock, and A. J. Read. 
1998. Autumn food habits of harbor porpoises, Phocoena 
pliocoena , in the Gulf of Maine. Fish Bull. 96:428-437. 
Gibson, R. N. 
1988. Development, morphometry and particle reten- 
tion capability of the gill rakers in the herring, Clupea 
harengus L. J. Fish Biol. 32:949-962. 
Greenstreet, S. P. R., J. A. McMillan, and E. Armstrong. 
1998. Seasonal variation in the importance of pelagic 
fish in the diet of piscivorous fish in the Moray Firth, 
