498 
Fishery Bulletin 107(4) 
(Mueter et al., 2002b). In addition, coastal SST during 
the first year at sea for sockeye salmon and within 400 
km of the point of ocean entry associated positively 
with survival rates of 19 stocks ranging from western 
Alaska to northern British Columbia and negatively with 
survival rates of 18 stocks in central British Columbia 
and Washington (Mueter et al., 2002a). Warmer winter 
SST (Nov-Feb) in offshore waters in the Gulf of Alaska 
were related to increases in total annual adult catch 
plus escapement of major B.C. and Alaska stocks (Pyper 
and Peterman, 1999). For the Karluk sockeye salmon, 
survival was correlated with spring coastal sea surface 
temperatures, winter and spring precipitation, and the 
more stormy winters before and following the juvenile 
stage, indicating two important marine periods in deter- 
mining the survival of Karluk sockeye salmon. This 
result is consistent with the hypotheses that climatic 
and oceanic conditions during the first few months at 
sea (Parker, 1971) and the first winter at sea (Beamish 
and Mahnken, 2001) are important in determining 
mortality rates. 
Climatic processes influencing salmon in the Pacific 
Northwest were not consistent with processes affecting 
Karluk sockeye salmon from Alaska. For example, water 
column mixing, as indicated by the Upwelling Index, 
did not correlate positively with faster growth or higher 
survival of Karluk sockeye salmon except in the winter 
and spring before the year of maturity. In the coastal 
waters off Oregon, by contrast, upwelling during the 
first year at sea of age-1 coho salmon was positively 
correlated with commercial catch of coho salmon from 
1947 to 1962 (Scarnecchia, 1981). The Northern Oscil- 
lation Index that is associated with a stronger trade 
winds, stronger coastal upwelling off Washington and 
Oregon, cooler surface and subsurface sea temperatures 
in waters off Washington, and higher Oregon hatchery 
salmon production (Schwing et al., 2002) was not cor- 
related with the growth and survival of Karluk sockeye 
salmon. 
The lack of a correlation between survival and 
growth of the age-2.2 sockeye salmon is similar to 
the findings of other studies where growth was de- 
termined from the scales of adult sockeye salmon. For 
example, Ml on the scales of adult sockeye salmon 
that returned to Egegik and Kvichak rivers in Bristol 
Bay, Alaska, was not correlated with variations in 
brood survival estimated from residuals of the Ricker 
model (Farley et al., 2007). From 1977 to 1997, growth 
on the scales of adult sockeye salmon from Bristol 
Bay showed reduced M2 and M3 growth and there 
was a smaller size-at-age for adults and lower smolt- 
to-adult survival rates for the even-year migrating 
smolts (Ruggerone et al., 2003). Alternatively, the 
size of juvenile sockeye salmon in Bristol Bay was 
positively correlated with survival rates of Bristol Bay 
sockeye salmon (Farley et al., 2007). The difference in 
results of correlating juvenile size with survival and 
with correlating adult scale growth with survival is 
likely due to size-selective mortality on the smaller 
fish during the juvenile stage (Farley et al., 2007; 
Table 4 
Correlation coefficients (r), P-values, and sample sizes 
in) relating survival to growth of sockeye salmon from 
Karluk River and Karluk Lake, Kodiak Island, Alaska. 
Survival is the residuals of the Ricker spawner-recruit- 
ment curve for Karluk sockeye salmon ( Oncorhynchus 
nerka). Smolt length (freshwater, FW), and juvenile (first- 
year marine, Ml), immature (second-year marine, M2), 
and maturing (third-year marine, M3) growth were esti- 
mated from measurement on the scales of age-2.2 sockeye 
salmon. No relationships between any pair of variables 
are significant at 5% (P<0. 0125 = 0. 05/4) indicated by *. 
Brood 
year 
Growth variable 
FW 
Ml 
M2 
M3 
1921-89 
0.073 
0.005 
0.136 
-0.290 
Survival 
r 
0.035 
0.315 
0.301 
-0.106 
p 
0.573 
0.967 
0.294 
0.022 
1948-89 
n 
62 
62 
62 
62 
Survival 
r 
0.035 
0.315 
0.301 
-0.106 
p 
0.835 
0.058 
0.070 
0.531 
n 
37 
37 
37 
37 
Moss et al., 2005) and a reduced detection of growth- 
survival relationships based on adult scales. 
Conclusion 
The first-, second-, and third-year marine scale growth, 
a proxy for the change in growth rate, of Karluk sock- 
eye salmon was not detected as a link between climate 
and survival for Karluk sockeye salmon. If the growth 
on the scales of adult sockeye salmon represents only 
a subset of the survivors (possibly fewer smaller and 
slower growing fish when mortality is high), then the 
weaker climate-growth-survival relationships recog- 
nized from adult scales provides insight into possible 
growth-climate-survival relationships. The growth- 
climate relationships and the mechanism for these 
relationships should further be assessed by relating 
the size and growth of juvenile salmon with physical 
and biological conditions measured at sea. 
Acknowledgments 
We thank the numerous agencies and individuals 
that collected information from sockeye salmon at the 
Karluk Research Station on Karluk River from 1924 
to 2000. Information sources included R. Gard and 
R. Bottorff, authors of a book on Karluk Lake, the 
National Archives in Anchorage, the National Marine 
Fisheries Service in Juneau, the Fisheries Research 
Institute at the University of Washington, and M. Wit- 
teveen, M. Foster, and P. Nelson at the Kodiak office 
for the Alaska Department of Fish and Game. 
