[1] Landcatch Nat Select Ltd, eCtr, Alloa FK10 3LP, Scotland
[2] Roslin Bioctr, Roslin Inst, Roslin EH25 9PS, Midlothian, Scotland
[3] Roslin Bioctr, Royal Dick Sch Vet Studies, Roslin EH25 9PS, Midlothian, Scotland
[4] Univ Edinburgh, Inst Evolutionary Biol, Sch Biol Sci, Ashworth Labs, Edinburgh EH9 3JT, Midlothian, Scotland