1. Department of Biology, University of North Carolina, Chapel Hill, NC 27599, USA
2. Mathematics and Biosciences Group, Max F. Perutz Laboratories and Faculty of Mathematics, University of Vienna, Nordbergstraße 15, A-1090 Vienna, Austria
3. Present address: Aix-Marseille University, UMR-CNRS 6632, Evolutionary Biology and Modeling, 3 Place Victor Hugo, 13331 Marseille, France