[1] Univ Oxford, Dept Zool, Oxford OX1 3PS, England
[2] Univ Exeter, Sch Biosci, Ctr Ecol & Conservat, Penryn TR10 9EZ, Cornwall, England
[3] Univ Birmingham, Sch Biosci, Birmingham B15 2TT, W Midlands, England
[4] Scottish Univ Environm Res Ctr, NERC, Life Sci Mass Spectrometry Facil, E Kilbride G75 0QF, Lanark, Scotland