1. Centre of Excellence in Evolutionary Research, Department of Biological and Environmental Science, P.O. Box 35, FI-40014, University of Jyväskylä, Finland
2. Natural History Museum, P.O. Box 35, FI-40014, University of Jyväskylä, Finland
3. Current address: ETH Zurich, Institute of Plant, Animal and Agroecosystem Sciences, Applied Entomology, Schmelzbergstrasse 9, CH-8092 Zurich, Switzerland