[1] Iowa State Univ, Dept Ecol Evolutionary & Organismal Biol, Ames, IA 50011 USA
[2] Univ Oulu, Ecol & Genet Res Unit, Oulu, Finland
[3] Norwegian Univ Sci & Technol, Dept Biol, Trondheim, Norway
[4] Norwegian Univ Sci & Technol, Grad Degree Program Ecol, Trondheim, Norway
[5] Colorado State Univ, Dept Biol, Ft Collins, CO USA