1. Department of Biology, University of Oxford , Oxford OX1 3RB , UK
2. The Roslin Institute, Royal (Dick) School of Veterinary Science, University of Edinburgh , Midlothian EH25 9RG , UK
3. Canada Research Chair in Forest Genomics, Forest Research Centre, Université Laval , Québec, QC GIV 0A6 , Canada
4. Institute for Systems and Integrative Biology, Université Laval , Québec, QC GIV 0A6 , Canada
5. Forest Research, Northern Research Station , Midlothian EH25 9SY , UK