1. The James Hutton Institute, Craigiebuckler, Aberdeen AB15 8QH, UK
2. Institute of Biological and Environmental Sciences, University of Aberdeen, Zoology Building, Tillydrone Avenue, Aberdeen AB24 2TZ, UK
3. Institute für Biologie—Botanik, Freie Universität Berlin, Altensteinstrasse 6, Berlin 14195, Germany
4. Evolution, Modeling and Analyses of Biodiversity group, Laboratoire d'Ecologie Alpine, UMR CNRS 5553, Université Joseph Fourier, Grenoble Cedex 9, France