[1] Heriot Watt Univ, Ctr Marine Biodivers & Biotechnol, Edinburgh EH14 4AS, Midlothian, Scotland
[2] Univ Glasgow, Sch Geog & Earth Sci, Glasgow G12 8QQ, Lanark, Scotland
[3] Plymouth Marine Lab, Plymouth PL1 3DH, Devon, England
[4] Univ N Carolina, Dept Biol Sci, Wilmington, NC 28403 USA
[5] Scottish Assoc Marine Sci, Oban PA37 1QA, Argyll, Scotland