[1] Plymouth Marine Lab, Marine Ecol & Biodivers, Prospect Pl, Plymouth PL1 3DH, Devon, England
[2] Univ Essex, Sch Biol Sci, Wivenhoe Pk, Colchester CO4 3SQ, Essex, England
[3] Dept Environm Food & Rural Affairs, Seacole Block,2 Marsham St, London SW1P 4DF, England
[4] Univ Plymouth, MBERC, Sch Biol & Marine Sci, Plymouth PL4 8AA, Devon, England