130 
Fishery Bulletin 99(1) 
Table 2 (continued) 
sMDH- 
sMDH- 
Total for 
Al,2 * 
Bl,2* 
mMEP-1* 
MPI 
PEPB* 
mAH-2* loci 
Collection site 
G 
df 
G 
df 
G 
df G 
df G 
df 
G 
df G 
df 
Monetka 
1.75 
4 
1.20 
6 
0.01 
2 
0.34 
2 
1.33 
2 
2.07 
1 
41.20 
59 
Ochepukha 
Other 
— 
25.79 
30 
3.28 
10 
— 
— 
— 
46.70 
90 
Znamenka 
0.11 
4 
37.29 
48 
4.14 
16 
1.35 
2 
8.47 
6 
0 
1 
121.04 
186 
Within Ochepukha 
0.11 
4 
63.08 
78 
7.43 
26 
1.35 
2 
8.47 
6 
0 
1 
167.74 
276 
Between Ochepukha 
0.03 
4 
6.60 
6 
1.25 
2 
1.87 
2 
3.16 
2 
— 
45.91 
58 
Total Ochepukha 
0.15 
8 
69.68 
84 
8.67 
28 
3.22 
4 
11.62 
8 
0 
1 
213.66 
334 
Within Sakhalin 
2.67 
16 
77.30 
114 
18.30 
38 
3.77 
8 
13.01 
12 
2.07 
2 
297.94 
496 
Among Sakhalin 
8.52 
12 
16.07 
18 
1.10 
6 
2.28 
6 
3.54 
6 
4.44 
3 
131.01 
177 
Total Sakhalin 
11.19 
28 
93.37 
132 
19.40 
44 
6.05 
14 
16.55 
18 
6.51 
5 
428.95 
673 
Northern Sea of Okhotsk 
0.60 
4 
0 
6 
0.66 
2 
0 
2 
9.55 h 
2 
— 
16.41 
41 
Western Kamchatka 
0.16 
12 
18.47 
24 
4.87 
8 
5.21 
8 
6.08 
8 
6.55 
3 
190.11 
220 
Eastern Kamchatka 
4.08 
4 
4.02 
6 
3.38 
2 
1.57 
2 
2.14 
2 
1.42 
1 
39.07 
61 
Within Asia 
16.11 
52 
118.60 
174 
29.07 
58 
12.82 
28 
40.96 
32 
18.78 
10 
723.41 
1057 
Among Asia 
74.76'' 
16 
45.82 fe 
24 
8.14 
8 
15.98° 
8 
76.91'' 
8 
15.69'’ 
4 
768.81'' 
236 
Total Asia 
90.87° 
68 
164.42 
198 
37.21 
66 
28.81 
36 
117.87'' 
40 
34.47' 1 
14 
1492.22'' 
1293 
North America 
Prince William Sound 
5.67 
16 
13.22 
24 
4.12 
8 
6.27 
8 
11.49 
8 
8.31° 
3 
176.78“ 
247 
Total within 
96.53 
84 
177.64 
222 
41.33 
74 
35.08 
44 
129.36 rf 
48 
42.78 
17 
1669.0H 
1540 
Between Alaska and Asia 
9.40 
4 
12.09 
6 
4.06 
2 
1.73 
2 
12.88 6 
2 
2.33 
1 
717.87'' 
62 
Total for collections 
105.93 
88 
189.73 
228 
45.39 
76 
36.81 
46 
142. 24 d 
50 
45.11' 
18 
2386.85'' 
1602 
"P < 0.05, h P < 0.01, P < 0.001, d P < 0.0001 
No overall heterogeneity was observed in tests among 
collections within each of the remaining geographic re- 
gions: northern Okhotsk coast, eastern Kamchatka, and 
western Kamchatka (Table 2). The northern Okhotsk and 
western Kamchatka collections each had a single locus 
that indicated heterogeneity, but neither test was signifi- 
cant after correction for multiple testing. 
The Alaska (Prince William Sound) collections exhibited 
low heterogeneity over all loci (G=176.78, 147 df; P=0.047) 
(Table 2); tests at three loci suggested that heterogeneity 
was not significant after correction for multiple testing. 
Although not the focus of this paper, the overall hetero- 
geneity suggests local genetic structure in even-year pink 
salmon within Prince William Sound. 
The collections of even-year pink salmon within the 
Asian geographic regions were relatively homogeneous, but 
we found strong heterogeneity among regions (G=1492.22, 
1257 df; PclCP 4 ) (Table 2). Of the nine loci that individu- 
ally suggest heterogeneity, eight ( GDA *, sAAT-3 *, PGDH *, 
PEPD-2 *, FH*, GR*, sMDH-Al *, and PEPB *) showed sig- 
nificant heterogeneity (P<0.05) after correction for multi- 
ple testing. The ratios of heterogeneities among regions 
to heterogeneity within regions (approximate P’s) for each 
locus (not shown) were significant (P<0.05) for 13 of 28 lo- 
ci, and comparisons of GDA*, sAAT-3 *, PGDH*, PEPD-2 *, 
FH*, GR*, sMDH-Al *, sMDH-B2* , and PEPB were high- 
ly significant (PcO.OOl). 
At the next level of hierarchy, between continents, Asian 
samples (in aggregate) differed overall from the Alaska 
samples with which they were compared (G=717.87, 62 df; 
P«10 -6 ) (Table 2); many of the loci examined contribute 
to the difference. After correction for multiple tests, GDA*, 
FDHG*, sAAT-3 *, PGDH*, TPI-2 *, in AH -4*, G3PDH-1*, 
GPI-B2*, LDH-A1*, and GR* were strongly significant 
(PcO.OOl). Of these loci, GDA*, sAAT-3 *, PGDH*, mAH-4 *, 
G3PDH-1* , LDH-A1 *, and GR* had appreciable variation 
(common allele <0.95) in at least some populations. These 
results suggest that at least seven allozyme loci may prove 
useful for distinguishing among even-year pink salmon 
stocks from different regions of the northern Pacific Ocean 
(Hawkins et ah, 1998). 
Data from 36 loci common to all regions were used 
to estimate pairwise chord distances (Cavalli-Sforza and 
Edwards, 1967) with which we constructed an unrooted 
neighbor-joining tree (Fig. 2). The tree supports a geo- 
graphic basis for the variability observed among collec- 
tions and suggests a geographic relationship among re- 
gions. Four clusters are evident along the tree axis: in 
linear order, they consisted of the collections from south- 
ern Okhotsk (Hokkaido Island and Sakhalin Island), west- 
ern Kamchatka, eastern Kamchatka, and Alaska. The 
greatest distance was between the Alaska cluster and all 
the other collections. When the Magadan samples were 
included in a similar tree with data from 34 loci, they 
