[1] Univ Groningen, Ctr Ecol & Evolutionary Studies, NL-9700 CC Groningen, Netherlands
[2] INRIA Res Team MERE, UMR Syst Anal & Biometr, F-34060 Montpellier, France
[3] ETH, Inst Integrat Biol, CH-8092 Zurich, Switzerland
[4] Cardiff Univ, Sch Earth & Ocean Sci, Cardiff CF10 3AT, S Glam, Wales
[5] Univ London Imperial Coll Sci Technol & Med, Div Biol, Ascot SL5 7PY, Berks, England