[1] GEOMAR Helmholtz Ctr Ocean Res Kiel, Wischhofstr 1-3, D-24148 Kiel, Germany
[2] Marine Biol Assoc UK, Citadel Hill, Plymouth PL1 2PB, Devon, England
[3] Univ Plymouth, Sch Biol & Marine Sci, Marine Biol & Ecol Res Ctr, Plymouth PL4 8AA, Devon, England