[1] Univ Cambridge, Dept Phys Dev & Neurosci, Cambridge, England
[2] Wellcome Trust Sanger Inst, Cambridge, England
[3] Univ Cambridge, Dept Genet, Cambridge CB2 3EH, England
[4] Univ Melbourne, Dept Zool, Melbourne, Vic 3010, Australia
[5] Australian Natl Univ, Res Sch Biol Sci, Canberra, ACT 2601, Australia