208 
Fishery Bulletin 117(3) 
absence of genetically distinct stock isolation. Examination 
of multiple biological metrics of Pacific sardine with testing 
of habitat prediction models should help elucidate migra¬ 
tion patterns and improve estimates of migration. A new 
perspective of greater interannual and seasonal variability 
in behavior of Pacific sardine will also likely benefit future 
research efforts and fisheries management. 
Acknowledgments 
We dedicate this manuscript to our inspirational coau¬ 
thor and good friend B. Emmett, who lost a battle against 
cancer prior to publication. We thank, for their assis¬ 
tance, individuals from the NOAA Northwest and South¬ 
west Fisheries Science Centers, Oregon State University, 
the Scripps Institution of Oceanography, Fisheries and 
Oceans Canada, and the Pacific sardine fishing industry. 
Funding was provided by the National Marine Fisheries 
Service and by the Mamie Markham Research and Walter 
G. Jones Fisheries Development Awards to R. Baldwin. 
Literature cited 
Anderson, M. J. 
2001. A new method for non-parametric multivariate analy¬ 
sis of variance. Austral Ecol. 26:32-46. 
Anderson, M. J., and J. Robinson. 
2003. Generalized discriminant analysis based on distances. 
Aust. N.Z. J. Stat. 45:301-318. 
Anderson, M. J., R. N. Gorley, and K. R. Clarke. 
2008. PERM AN OVA+ for PRIMER: guide to software and 
statistical methods, 214 p. PRIMER-e, Plymouth, UK. 
Arthur, J. R., and E. Albert. 
1994. A survey of the parasites of Greenland halibut (Rein- 
hardtius hippoglossoides ) caught off Atlantic Canada, 
with notes on their zoogeography in this fish. Can. J. Zool. 
72:765-778. 
Bakun, A. 
1996. Patterns in the ocean: ocean processes and marine pop¬ 
ulation dynamics, 323 p. Calif. Sea Grant Coll. Syst., Univ. 
Calif., La Jolla, CA. 
Baldwin, R. E., and C. P. Goater. 
2003. Circulation of parasites among fishes from lakes in 
the Caribou Mountains, Alberta, Canada. J. Parasitol. 
89:215-225. 
Baldwin, R. E., T. W. Miller, R. D. Brodeur, and K. C. Jacobson. 
2008. Expanding the foraging history of juvenile Pacific 
salmon: combining stomach-content and macroparasite- 
community analyses for studying marine diets. J. Fish Biol. 
72:1268-1294. 
Baldwin, R. E., M. B. Rew, M. L. Johansson, M. A. Banks, and 
K. C. Jacobson. 
2011. Population structure of three species of Anisakis nem¬ 
atodes recovered from Pacific sardines (Sardinops sagax) 
distributed throughout the California Current System. 
J. Parasitol. 97:545-554. 
Baldwin, R. E., M. A. Banks, and K. C. Jacobson. 
2012. Integrating fish and parasite data as a holistic 
solution for identifying the elusive stock structure of 
Pacific sardines ( Sardinops sagax). Rev. Fish Biol. Fish. 
22:137-156. 
Bentley, P. J., R. L. Emmett, N. C. H. Lo, and H. G. Moser. 
1996. Egg production of Pacific sardine ( Sardinops sagax) off 
Oregon in 1994. CalCOFI Rep. 37:193-200. 
Blaylock, R. B., L. Margolis, and J. C. Holmes. 
1998. Zoogeography of the parasites of Pacific halibut ( Hip- 
poglossus stenolepis) in the northeast Pacific. Can. J. Zool. 
76:2262-2273. 
2003. The use of parasites in discriminating stocks of the 
Pacific halibut ( Hippoglossus stenolepis) in the northeast 
Pacific. Fish. Bull. 101:1-9. 
Bray, J. R., and J. T. Curtis. 
1957. An ordination of the upland forest communities of 
southern Wisconsin. Ecol. Monogr. 27:325-349. 
Bush, A. O., K. D. Lafferty, J. M. Lotz, and A. W. Shostak. 
1997. Parasitology meets ecology on its own terms: Margolis 
et al. revisited. J. Parasitol. 83:575-583. 
Calambokidis, J., and J. Barlow. 
2004. Abundance of blue and humpback whales in the east¬ 
ern North Pacific estimated by capture-recapture and 
line-transect methods. Mar. Mamm. Sci. 20:63-85. 
Catalano, S. R., I. D. Whittington, S. C. Donnellan, and B. M. 
Gillanders. 
2014. Parasites as biological tags to assess host population 
structure: guidelines, recent genetic advances and com¬ 
ments on a holistic approach. Int. J. Parasitol: Parasites 
Wildl. 3:220-226. 
Clark, F. N., and J. F. Janssen. 
1945. Movements and abundance of the sardine as mea¬ 
sured by tag returns. In Results of tagging experiments 
in California waters on the sardine (Sardinops caerulea). 
Calif. Dep. Fish Game, Fish Bull. 61, p. 7-42. [Available 
from website]. 
Clarke, K. R., and R. H. Green. 
1988. Statistical design and analysis for a ‘biological effects’ 
study. Mar. Ecol. Prog. Ser. 46:213-226. 
Clarke, K. R., and R. N. Gorley. 
2015. PRIMER v7: user manual/tutorial, 296 p. PRIMER-e, 
Plymouth, UK. 
Clarke, K. R., P. J. Somerfield, and M. G. Chapman. 
2006. On resemblance measures for ecological studies, 
including taxonomic dissimilarities and a zero-adjusted 
Bray-Curtis coefficient for denuded assemblages. J. Exp. 
Mar. Biol. Ecol. 330:55-80. 
Clarke, K. R., R. N. Gorley, P. J. Somerfield, and R. M. Warwick. 
2014a. Change in marine communities: an approach to statis¬ 
tical analysis and interpretation, 3 rd ed., 260 p. PRIMER-e, 
Plymouth, UK. 
Clarke, K. R., J. R. Tweedley, and F. J. Valesini. 
2014b. Simple shade plots aid better long-term choices of 
data pre-treatment in multivariate assemblage studies. 
J. Mar. Biol. Assoc. U.K. 94:1-16. 
Cury, P, A. Bakun, R. J. M. Crawford, A. Jarre, R. A. Quinones, 
L. J. Shannon, and H. M. Verheye. 
2000. Small pelagics in upwelling systems: patterns of inter¬ 
action and structural changes in “wasp-waist” ecosystems. 
ICES J. Mar. Sci. 57:603-618. 
de Moor, C. L., D. S. Butterworth, and C. D. van der Lingen. 
2017. The quantitative use of parasite data in multistock 
modelling of South African sardine (Sardinops sagax). Can. 
J. Fish. Aquat. Sci. 74:1895-1903. 
Demer, D. A., J. P. Zwolinski, K. A. Byers, G. R. Cutter, J. S. Ren- 
free, T. S. Sessions, and B. J. Macewicz. 
2012. Prediction and confirmation of seasonal migration of 
Pacific sardine (Sardinops sagax) in the California Current 
Ecosystem. Fish. Bull. 110:52-70. 
