[1] Univ Edinburgh, Roslin Inst, Edinburgh Genom, Edinburgh EH25 9RG, Midlothian, Scotland
[2] Univ Edinburgh, RD SVS, Edinburgh EH25 9RG, Midlothian, Scotland
[3] Univ Edinburgh, Edinburgh Genom, Inst Evolutionary Biol, Ashworth Labs, Edinburgh EH9 3JT, Midlothian, Scotland