[1] Univ Oslo, Dept Biol, Div Zool, N-0316 Oslo, Norway
[2] Univ Copenhagen, Inst Zool, Dept Populat Ecol, DK-1200 Copenhagen, Denmark
[3] Abisko Sci Res Stn, S-98107 Abisko, Sweden
[4] Univ Sheffield, Dept Anim & Plant Sci, Sheffield Ctr Arctic Ecol, Sheffield S10 2BR, S Yorkshire, England