1. Department of Biology, University of Copenhagen, Ole Maaløes Vej 5, DK-2200 Copenhagen N, Denmark; 2Instituto Gulbenkian de Ciência, Rua da Quinta Grande 6, P-2780-156 Oeiras, Portugal; 3Department of Integrative Biology, University of California Berkeley, 1005 Valley Life Sciences Building, Berkeley, CA 94720, USA; and 4Centre for GeoGenetics, Natural History Museum, University of Copenhagen, Øster Voldgade 5–7, DK-1350 Copenhagen K, Denmark