[1] Univ Tecn Lisboa, Ctr Estudos Florestais, P-1349017 Lisbon, Portugal
[2] Univ Tecn Lisboa, Dept Matemat, Inst Super Agron, P-1349017 Lisbon, Portugal
[3] Univ Tecn Lisboa, CEMAPRE, P-1200781 Lisbon, Portugal
[4] Univ Tecn Lisboa, Inst Sup Econ & Gestao, P-1200781 Lisbon, Portugal
[5] Univ Helsinki, Dept Biol & Environm Sci, Metapopulat Res Grp, FIN-00014 Helsinki, Finland
[6] CSIC, Natl Museum Nat Sci, Dept Biodivers & Evolutionary Biol, ES-28006 Madrid, Spain
[7] Univ Sheffield, Dept Anim & Plant Sci, Sheffield S10 2TN, S Yorkshire, England