[1] Plymouth Marine Lab, Plymouth PL1 3DH, Devon, England
[2] Danish Inst Fisheries & Marine Res, Dept Ecol & Aquaculture, DK-2920 Charlottenlund, Denmark
[3] Univ Gothenburg, Dept Marine Ecol, Kristianberg Marine Res Stn, S-45034 Fiskebackskil, Sweden