Crow et a!.: Vertical distribution, diet, and reproduction of Zameus squamulosus 
21 1 
Table 2 
The dietary contents found in the stomachs (S) and valvular intestines (VI) of female velvet dog¬ 
fish ( Zameus squamulosus ) captured in Hawaii longline fisheries between 1995 and 2008. Nine 
of the 21 sharks examined had recognizable dietary contents. 
No. of S and VI 
Total number of 
with dietary items 
dietary items 
Cephalopod 
Squid 
7 
20 
Nototodarus hawaiiensis 
1 
1 
Sandalops melancholicus 
1 
1 
Histioteuthis sp. 
2 
4 
Onykia sp. 
1 
1 
Walvisteuthis sp. 
1 
1 
Unidentified squid 
4 
12 
Crustacean 
Unidentified shrimp 
1 
1 
Fish 
Unidentified shark 
1 
1 
Unidentified teleost 
4 
4 
Table 3 
Number of female velvet dogfish (Zameus squamulosus), collected and retained each month from 
Hawaii longline fisheries between 1995 and 2008, classified as immature (IM) or in 1 of 7 stages 
of reproductive maturity: nongravid females with developing ovarian 
eggs (DO), nongravid fe- 
males holding at least one large ovarian egg (LO), ovulating females (OV), postovulatory females 
with uterine 
ova (UO), gravid females carrying at least one encapsulated developing embryo (DE), 
gravid females carrying near-term embryos free in the uterus (NE), and postpartum females (PP). 
Table format 
is adapted from Tanaka et al. (1990) and Yano (1995). 
Month 
IM DO LO 
OV UO DE 
NE PP Total 
Jan 
0 
Feb 
1 
1 
Mar 
1 
1 
Apr 
1 
1 
2 
May 
1 
1 
Jun 
0 
Jul 
2 
1 3 
Aug 
1 
1 2 
Sep 
1 
1 
2 
Oct 
2 
2 
Nov 
1 1 
2 
4 
Dec 
1 
1 
2006; Zhu et al., 2012; Romanov et al., 2013) indicate 
that female velvet dogfish clearly exhibit an epipelagic 
aspect in their behavior. The capture of this species at 
depths as great as 2000 m also reveals a large vertical 
and benthic range in their movements (Pajuelo et al., 
2010; Romanov et al., 2013). In general, larger squa- 
loid sharks similar in size to velvet dogfish tend to be 
benthic, slope, or shelf dwellers, whereas the majority 
of oceanic dogfish species are smaller species (Hubbs 
et al., 1967; Pikitch et al., 2008). Female velvet dogfish 
regularly inhabit pelagic waters, and this relatively 
large squaloid may transition from bottom to midwater 
or pelagic waters to take advantage of feeding oppor¬ 
tunities, probably by following migrations of midwater 
squid and fish (Parin, 1970; Krefft, 1980). 
The capture of only females off Hawaii raises the 
possibility of sexual segregation, because females more 
frequently use the epipelagic environment and males 
remain more bottom oriented at greater depths. Sexual 
segregation for this species at other locations is indi¬ 
cated by the capture of only females on floating long- 
lines in the northwestern North Pacific Ocean (Yano 
