[1] Univ Gothenburg, Dept Marine Sci, Box 461, SE-40530 Gothenburg, Sweden
[2] Univ Gothenburg, Dept Biol & Environm Sci, Box 461, SE-40530 Gothenburg, Sweden
[3] Univ Calif San Diego, Scripps Inst Oceanog, 9500 Gilman Dr, La Jolla, CA 92093 USA
[4] Univ Copenhagen, Marine Biol Sect, Dept Biol, Strandpromenaden 5, DK-3000 Helsingor, Denmark
[5] Inst Marine Res, Fram Ctr, N-9007 Tromso, Norway