1. Ecology and Evolutionary Biology, School of Biosciences University of Sheffield Western Bank Sheffield S10 2TN UK
2. Laboratoire Evolution et Diversité Biologique UMR5174, CNRS/IRD/Université Toulouse 3 Toulouse 31062 France
3. Department of Biology University of Fribourg Chemin du Musée 10 Fribourg 1700 Switzerland