210 
Fishery Bulletin 116(2) 
Table 1 
Capture records and reproductive biology for female velvet dogfish (Zameus squamulosus ) collected from Hawaii longline 
fisheries between 1995 and 2008. The letters a and b indicate different velvet dogfish caught on the same longline set. 
N/A=not available. 
Total Estimated 
Capture length hook depth Maturity 
ID no. date (mm) (m) status Reproductive status 
1 
15 Oct 95 
786 
30 
Mature 
2 
N/A 
746 
N/A 
Mature 
3 
24 Sep 01 
720 
N/A 
Mature 
4 
07 Oct 01 
735 
24 
Mature 
5 
12 Nov 01 
758 
400 
Mature 
6 
10 Nov 01 
594 
305 
Immature 
7 
16 Dec 01 
728 
250 
Immature 
8 
10 Feb 02 
576 
364 
Immature 
9 
N/A 
728 
N/A 
Mature 
10 
28 Jul 02 
752 
110 
Mature 
11 
26 Aug 02 
791 
180 
Mature 
12 
10 Sep 02 
729 
300 
Immature 
13 
24 Mar 03 
736 
120 
Mature 
14 
23 Nov 08 
805 
N/A 
Mature 
15 
08 May 08 
715 
83 
Mature 
16 
27 Apr 08 
720 
71 
Mature 
17 
18 Aug 06 
751 
182 
Mature 
18 
28 Apr 08 
722 
82 
Mature 
19 
12 Nov 01 
720 
400 
Mature 
20 a 
07 Jul 06 
742 
182 
Mature 
21 b 
07 Jul 06 
839 
182 
Mature 
8 ovarian eggs; largest 37 mm; uterus width 18.5 mm 
7 ovarian eggs; largest 52.8 mm 
Uterine eggs; largest 47.9 mm 
6 ovarian eggs; range 35.5-47.5 mm 
9 ovarian eggs; range 44.4-56.5 mm 
No sexual development 
No sexual development 
No sexual development 
Uterine eggs not measured 
Small ovarian eggs: largest 8.2 mm; uterus width 29-33 mm 
4 embryos; range 31.4-36.7 mm TL 
9 ovarian eggs: largest 34.4 mm; uterus width 8.6 mm 
9 ovarian eggs; range 18.9-22.3 mm 
8 uterine eggs (not measured) 
8 embryos; range 44.5-60.4 mm TL 
Ovary starting to develop; uterus width 10.5 mm 
Ovary and uteri developed 
Eggs ruptured in abdominal cavity 
10 ovarian eggs; range 25.3-28.8 mm; uterus width 15.5 mm 
10 uterine eggs (not measured); uterus width 41.1 mm 
9 uterine eggs; range 38.5-47.7 mm 
was present in 1 shark. Identifiable squid beaks rep¬ 
resented 4 families; 2 identified species, Nototodarus 
hawaiiensis and Sandalops melancholicus; and 3 rec¬ 
ognizable genera, Histioteuthis, Onykia, and Walvisteu- 
this (Table 2). 
Female reproductive biology 
Seventeen of the 21 examined velvet dogfish were ma¬ 
ture. Both left and right ovaries of female velvet dog¬ 
fish were functional. Uteri were noncompartmentalized 
and contained 6-10 ova that were between 36.7 and 
52.8 mm in diameter. It is unclear whether all uter¬ 
ine eggs were fertilized. Four females between 576 and 
729 mm TL had small, developing ovaries (< 10 mm in 
diameter) and narrow uteri (<10 mm in width) and 
were classified as immature (Table 1). Seventeen fe¬ 
males, ranging in size between 715 and 839 mm TL, 
had ova >10 mm in diameter and expanded uteri (> 10 
mm in width) and were classified as mature (Table 1). 
Six of the 7 stages of reproductive maturity for females 
were observed in the specimens from Hawaii. No near- 
term embryos were found. Large ova (>30 mm) were 
observed in ovaries of velvet dogfish captured during 
September, October, and November. The smallest preg¬ 
nant female was 715 mm TL. Uterine eggs were ob¬ 
served in specimens collected during July, September, 
and November, and developing embryos were found in 
uteri of sharks obtained in May and August (Table 3). 
No reproductive seasonality could be detected from our 
limited sample size (Tables 1 and 3). 
Brood sizes of 2 pregnant females were 4 and 8 em¬ 
bryos, and these ranged from 31.4 to 36.7 mm TL in 
August and between 45.5 and 60.4 mm TL in May. On 
the basis of the presence of yolk sacs and size of the 
embryos, we determined that none of them were con¬ 
sidered near-term embryos. 
Discussion 
Vertical distribution 
The capture of velvet dogfish in Hawaii pelagic long- 
line fisheries, is not as rare an event as previously 
suggested (Wetherbee and Crow, 1996). The worldwide 
distribution of this species includes a number of in¬ 
sular locations, such as Hawaii, and other waters in 
the Pacific Ocean, the Atlantic Ocean, and the Indian 
Ocean, and indicates that this species is associated 
with islands in addition to continental slope habitats. 
The estimated hooking depths for shark captures off 
Hawaii (24-400 m) and similar pelagic depths reported 
in other studies (Yano and Tanaka, 1984; Dai et ah, 
