[1] Univ Helsinki, Fac Biol & Environm Sci, Res Ctr Ecol Change, Organismal & Evolutionary Biol Res Programme, Helsinki, Finland
[2] Univ Massachusetts Amherst, Dept Environm Conservat, Amherst, MA USA
[3] Nat Resources Inst Finland Luke, Helsinki, Finland
[4] Univ Helsinki, Fac Sci, Dept Math & Stat, Helsinki, Finland