[1] Univ Bristol, Sch Biol Sci, Bristol BS8 1TQ, England
[2] Univ Oxford, Dept Biol, Oxford OX1 3RB, England
[3] Univ Catania, Dept Biol Geol & Environm Sci, I-95128 Catania, Italy
[4] Univ Naples Federico II, Dept Biol, I-80126 Naples, Italy
[5] UCL, Dept Genet Evolut & Environm, London WC1E 6BT, England
[6] Monash Univ, Sch Biol Sci, Melbourne, Vic 3800, Australia