[1] Plymouth Marine Lab, Plymouth PL1 3DH, Devon, England
[2] Akvaplan Niva AS, High North Res Ctr Climate & Environm, N-9296 Tromso, Norway
[3] Univ Exeter, Coll Engn Math & Phys Sci Phys, Exeter EX4 4QL, Devon, England
[4] Univ Exeter, Coll Life & Environm Sci Biosci, Exeter EX4 4QD, Devon, England