1. Mathematical Institute, Leiden University, 2333CA Leiden, The Netherlands
2. Institute of Biology, Leiden University, 2333CA Leiden, The Netherlands
3. Evolution and Ecology Program, International Institute for Applied Systems Analysis, 2361 Laxenburg, Austria
4. Naturalis Biodiversity Center, 2333CR Leiden, The Netherlands
5. Department of Medical Genetics, Universitair Medisch Centrum Utrecht, Sectie Complex Genetics, Utrecht, The Netherlands