1. Institute of Evolutionary Biology , University of Edinburgh , King’s Buildings, West Mains Road, Edinburgh , EH9 3JT , United Kingdom of Great Britain and Northern Ireland .
2. Forestry Research , Northern Research Station , Roslin, Midlothian, EH25 9SY , United Kingdom of Great Britain and Northern Ireland .
3. Rothamsted Research , Harpenden, Hertfordshire, AL5 2JQ , United Kingdom of Great Britain and Northern Ireland .
4. The Roslin Institute, Royal(Dick) School of Veterinary Studies , University of Edinburgh , Roslin Biocentre, Midlothian, EH25 9PS , United Kingdom of Great Britain and Northern Ireland .