300 
Fishery Bulletin 111(4) 
Table 2 
Results of permutational analysis of variance (PERMANOVA) with fork length data from our surveys of 4 species — 
Opakapaka (Pristipomoides filamentosus ), Kalekale (P. sieboldii), Onaga ( Etelis coruscans), and Ehu (E. carbuncu- 
lus ) — in the main Hawaiian Islands between May 2007 and June 2009. The following factors were tested within the 
preferred depths of each species: bottomfish restricted fishing area location (BR), protection (PR), and habitat type 
(HA). Preferred depths are noted in the column head for each species. df=degrees of freedom; P=PERMANOVA F- 
statistic; P=PERMANOVA P-value. Asterisks indicate statistical significance at P<0.05. 
Factor 
Opakapaka 
(90-210 m) 
Kalekale 
(180-270 m) 
Onaga 
(210-300 m) 
Ehu 
(210-300 m) 
df 
F 
P 
df 
F 
P 
df 
F 
P 
df 
F 
P 
BR 
5 
36.04 
0.00* 
5 
28.20 
0.00* 
4 
11.05 
0.00* 
4 
4.90 
0.00* 
PR 
1 
14.24 
0.00* 
1 
1.43 
0.23 
0 
No test 
0 
No test 
HA 
3 
11.39 
0.00* 
3 
18.38 
0.00* 
1 
0.48 
0.49 
2 
1.77 
0.17 
BRxPR 
5 
2.02 
0.08 
3 
16.57 
0.00* 
3 
4.82 
0.00* 
4 
0.84 
0.52 
BRxHA 
9 
7.66 
0.00* 
4 
1.16 
0.33 
1 
23.69 
0.00* 
5 
1.62 
0.16 
PRxHA 
2 
0.45 
0.64 
2 
0.21 
0.82 
2 
0.48 
0.61 
2 
1.31 
0.27 
BRxPRxHA 
5 
3.42 
0.01* 
2 
0.21 
0.81 
1 
13.26 
0.00* 
ONo test 
Residual 
419 
446 
242 
274 
for Ehu. The highest relative abundance for this spe- 
cies was in Pailolo Channel, and the lowest levels were 
seen at Niihau, Kaena, and Makapuu (Fig. 3; post hoc 
PERMANOVA, P<0.05). Overall, hard-low habitats had 
significantly greater numbers of Ehu than did other 
habitat types. By BRFA location and habitat type, the 
mean MaxNo of Ehu in Pailolo Channel was higher 
for hard-low than for soft-low habitats, and similar 
abundance estimates were found for hard-high, hard- 
low, and soft-high habitats on Penguin Bank. Niihau 
and Kaena differed from the other sampled locations 
in that hard-high habitats had a greater relative abun- 
dance of Ehu than did hard-low habitats. 
In our evaluation of mean lengths, BRFA location, 
protection, and habitat type were all important factors, 
and the interactions between them were sometimes 
significant (Table 2). BRFA location, protection, habitat 
type, the interaction of BRFA location and habitat type, 
and the interaction of all 3 factors were significant for 
Opakapaka. Niihau had the largest Opakapaka on 
average (65.29 cm FL) among sampled locations, and 
the smallest Opakapaka (28.35 cm FL; Fig. 4; post hoc 
PERMANOVA, P<0.05) were seen at Hilo. The smallest 
individual at Hilo measured ~16 cm FL, and the largest 
individual at Niihau was ~79 cm FL. Opakapaka from 
outside protected areas had a mean length of 42.89 cm 
FL and were larger than those fish observed inside the 
sampled BRFAs (40.53 cm FL; PERMANOVA, P<0.05). 
The smallest mean lengths of this species were found 
over hard-low habitats compared with other habitat 
types overall, other habitats at each BRFA location, 
and other habitats either inside or outside a particular 
BRFA (Fig. 4; Table 3; post hoc PERMANOVA, P<0.05). 
BRFA location, habitat type, and the interaction of 
BRFA location and protection were significant for Kale- 
kale. Pair-wise comparisons showed that this species 
had its smallest mean length (23.64 cm FL) at Kaena, 
was largest in hard-high habitats (31.46 cm FL) and 
smallest in soft-low habitats (8.64 cm FL, n= 2), and 
was larger inside the Penguin Bank and Pailolo Chan- 
nel BRFAs and outside the Hilo BRFA than in other 
sampled areas (Fig. 4; Table 3; post hoc PERMANOVA, 
P<0.05). The smallest individual Kalekale, however, 
measured 7.63 cm FL at Niihau. BRFA location, the 
interaction of BRFA location with protection, the in- 
teraction of BRFA location with habitat type, and the 
interaction of all 3 of these factors were significant for 
Onaga. Mean length for Onaga was smallest in Pailolo 
Channel (42.80 cm FL) than at other locations (Fig. 4) 
but larger inside the Pailolo Channel BRFA than out- 
side this protected area (Table 3; post hoc PERMANO- 
VA, P<0.05). The smallest individual Onaga measured 
15.05 cm FL. Although the interaction of BRFA loca- 
tion and habitat type and the interaction of BRFA 
location, protection, and habitat type had significant 
results for Onaga, no clear trends were seen. BRFA 
location was the only factor that had an influence on 
mean length for Ehu (Table 2; PERMANOVA, P< 0.05). 
Overall, mean sizes were very similar for this species 
but were smallest at Makapuu and Hilo (Fig. 4). 
For all sampled locations combined, size-related 
shifts in species-habitat associations were evident. 
The linear regressions of FL against depth for each 
species showed that size increased with depth for 
Opakapaka (coefficient of determination [r 2 ] =0.438, 
PcO.Ol) but did not for the other 3 species (Fig. 5). In 
our evaluation of the proportion of fish measured in 
each habitat type by size class, habitat associations 
clearly varied by size for Opakapaka, Kalekale, and 
Onaga (Fig. 6). Ehu had very similar habitat associa- 
