[1] Univ Bristol, Sch Biol Sci, Life Sci Bldg,24 Tyndall Ave, Bristol BS8 1TQ, Avon, England
[2] Univ Bristol, Sch Social & Community Med, Oakfield House, Bristol BS8 2BN, Avon, England
[3] Mem Univ Newfoundland, Dept Biol, St John, NF A1B 3X9, Canada
[4] Mem Univ Newfoundland, Dept Math & Stat, St John, NF A1C 5S7, Canada
[5] Cardiff Univ, Sch Biosci, Cardiff CF10 3AX, S Glam, Wales
[6] UCL Genet Inst, Gower St, London WC1E 6BT, England
[7] Univ Nebraska Lincoln, Sch Biol Sci, 424 Manter Hall,1104 T St, Lincoln, NE 68588 USA