[1] Univ Copenhagen, Zool Museum, DK-2100 Copenhagen, Denmark
[2] Univ Copenhagen, Ctr Macroecol, Dept Biol, DK-2100 Copenhagen, Denmark
[3] Princeton Univ, Dept Ecol & Evolut Biol, Princeton, NJ 08544 USA
[4] Univ Leeds, Inst Integrat & Comparat Biol, Leeds LS2 9JT, W Yorkshire, England
[5] Univ Washington, Dept Biol, Seattle, WA 98195 USA