[1] Sir Alister Hardy Fdn Ocean Sci, Lab, Plymouth PL1 2BN, Devon, England
[2] Ctr Environm Fisheries & Aquaculture Sci, Lowestoft, Suffolk, England
[3] Finnish Inst Marine Res, FIN-00931 Helsinki, Finland
[4] Univ Coll Swansea, Dept Biol Sci, Swansea, W Glam, Wales