[1] Univ New South Wales, Evolut & Ecol Res Ctr, Sydney, NSW, Australia
[2] Univ New South Wales, Sch Biol Earth & Environm Sci, Sydney, NSW, Australia
[3] Univ New South Wales, Sch Math & Stat, Sydney, Australia
[4] Australian Natl Univ, Res Sch Biol, Div Ecol & Evolut, Canberra, ACT, Australia
[5] Zhejiang Univ, Dept Biosyst Engn, Hangzhou, Peoples R China
[6] Univ Alberta, Dept Biol Sci, Edmonton, AB, Canada
[7] Okinawa Inst Sci & Technol Grad Univ, Theoret Sci Visiting Program, Onna, Japan