[1] Univ Sheffield, Dept Anim & Plant Sci, Western Bank, Sheffield S10 2TN, S Yorkshire, England
[2] Norwegian Inst Nat Res, Polar Environm Ctr, NO-9296 Tromso, Norway
[3] Marine Biol Lab, Ctr Ecosyst, Woods Hole, MA 02543 USA
[4] Royal Swedish Acad Sci, Abisko Sci Res Stn, S-98107 Abisko, Sweden