1. Centre for Applied Entomology and Parasitology, Keele University,Staffordshire ST5 5BG, UK
2. School of Biological Sciences, University of Manchester, 2.205 Stopford Building, Oxford Road, Manchester M13 9PT, UK
3. School of Life Sciences, Keele University, Staffordshire ST5 5BG,UK
4. Author for correspondence (e-mail: h.hurd@keele.ac.uk)