1. School of Natural and Environmental Sciences Newcastle University Newcastle upon Tyne NE1 7RU UK
2. Centre for Ecology and Hydrology Maclean Building, Benson Lane, Crowmarsh Gifford Wallingford Oxfordshire OX10 8BB UK
3. Butterfly Conservation Manor Yard, East Lulworth Wareham Dorset BH20 5QP UK
4. School of Biological, Biomedical and Environmental Sciences University of Hull Cottingham Road Hull HU6 7RX UK