[1] Newcastle Univ, Sch Marine Sci & Technol, Newcastle Upon Tyne NE1 7RU, Tyne & Wear, England
[2] Rhodes Univ, Dept Ichthyol & Fisheries Sci, ZA-6140 Grahamstown, South Africa
[3] Newcastle Univ, Sch Civil Engn & Geosci, Newcastle Upon Tyne NE1 7RU, Tyne & Wear, England
[4] Univ Edinburgh, Inst Quantitat Biol Biochem & Biotechnol, Sch Biol Sci, Edinburgh EH9 3FF, Midlothian, Scotland