[1] Umea Univ, Dept Math & Math Stat, Integrated Sci Lab, SE-90187 Umea, Sweden
[2] Int Inst Appl Syst Anal IIASA, Evolut & Ecol Program, Schlosspl 1, A-2361 Laxenburg, Austria
[3] Grad Univ Adv Studies Sokendai, Dept Evolutionary Studies Biosyst, Hayama, Kanagawa 2400193, Japan
[4] Stellenbosch Univ, Ctr Invas Biol, Dept Math Sci, ZA-7602 Matieland, South Africa
[5] African Inst Math Sci, Math & Phys Biosci, ZA-7945 Cape Town, South Africa