[1] UCL, Dept Genet Evolut & Environm, Gower St, London WC1E 6BT, England
[2] Zool Soc London, Inst Zool, Regents Pk, London NW1 4RY, England
[3] Tanzania Fisheries Res Inst, POB 9750, Dar Es Salaam, Tanzania
[4] UCL, Ctr Biodivers & Environm Res, Gower St, London WC1E 6BT, England