1. School of Natural Sciences, Bangor University, Deiniol Road, Bangor, Gwynedd, LL57 2UW, UK
2. Present address: Birmingham Institute of Forest Research, Institute of Microbiology and Infection, School of Biosciences, University of Birmingham, Birmingham, B15 2TT, UK
3. Forest Research, Centre for Forestry and Climate Change, Alice Holt Lodge, Farnham, Surrey GU10 4LH, UK