[1] Univ Aberdeen, Oceanlab, Newburgh AB41 6AA, Aberdeen, Scotland
[2] Univ Calif San Diego, Scripps Inst Oceanog, Ctr Marine Biodivers & Conservat, La Jolla, CA 92093 USA
[3] Univ Calif San Diego, Scripps Inst Oceanog, Integrat Oceanog Div, La Jolla, CA 92093 USA
[4] Japan Agcy Marine Earth Sci & Technol, Yokosuka, Kanagawa 2370061, Japan