[1] Univ Jyvaskyla, Dept Biol & Environm Sci, FIN-40351 Jyvaskyla, Finland
[2] Univ Helsinki, Div Populat Biol, Integrat Ecol Unit, FIN-00014 Helsinki, Finland
[3] Lund Univ, Dept Theoret Ecol, S-22362 Lund, Sweden
[4] Aalto Univ, Syst Anal Lab, FIN-02015 Helsinki, Finland