[1] Univ Siena, Dept Phys Earth & Environm Sci, Global Ocean Team, Nat Conservancy, Plan dei Mantellini 44, I-53100 Siena, Italy
[2] Univ Cambridge, Dept Zool, Cambridge CB2 3EJ, England
[3] Newcastle Univ, Sch Biol, Newcastle Upon Tyne NE1 7RU, Tyne & Wear, England
[4] Univ Highlands & Isl, Environm Res Inst, Thurso KW14 7EE, Scotland