[1] Natl Taiwan Ocean Univ, Dept Environm Biol & Fisheries Sci, Keelung 20224, Taiwan
[2] Natl Taiwan Ocean Univ, George Chen Shark Res Ctr, Keelung 20224, Taiwan
[3] Moss Landing Marine Labs, Pacific Shark Res Ctr, Moss Landing, CA 95039 USA
[4] Calif Acad Sci, Dept Ichthyol, San Francisco, CA 94118 USA
[5] South African Inst Aquat Biodivers, ZA-6140 Grahamstown, South Africa