1 Lancaster Environment Centre, Lancaster University, Lancaster, LA1 4YQ, UK.
2 Department of Zoology, Edward Grey Institute, University of Oxford, Oxford, OX1 3PS, UK.
3 CNRS, UMR 5558 "Biométrie et Biologie Evolutive", Université de Lyon, Université Lyon 1, Villeurbanne, France.
4 Center for Macroecology, Evolution and Climate Change, Natural History Museum of Denmark, University of Copenhagen, Universitetsparken 15, København, Denmark.