[1] British Antarctic Survey, Nat Environm Res Council, Cambridge CB3 0ET, England
[2] Univ Exeter, Ctr Ecol & Conservat, Penryn Campus, Penryn TR10 9EZ, Cornwall, England
[3] Univ Coimbra, Dept Life Sci, Marine & Environm Sci Ctr MARE UC, P-3000456 Coimbra, Portugal
[4] La Rochelle Univ, CNRS, UMR 7372, CEBC, F-79360 Villiers En Bois, France