[1] Univ Aberdeen, Oceanlab, Sch Biol Sci, Aberdeen AB41 6AA, Scotland
[2] Heriot Watt Univ, Ctr Marine Biodivers & Biotechnol, Sch Life Sci, Edinburgh EH14 4AS, Midlothian, Scotland
[3] Univ N Carolina, Ctr Marine Sci, 601 S Coll Rd, Wilmington, NC 28403 USA
[4] Scottish Marine Inst, Scottish Assoc Marine Sci, Oban PA37 1QA, Argyll, Scotland