[1] Univ Bristol, Life Sci Bldg, Bristol BS8 1TQ, Avon, England
[2] Univ Oxford, Dept Zool, Oxford OX1 3PS, England
[3] Univ Bristol, Dept Comp Sci, Bristol BS8 1UB, Avon, England
[4] Univ Bordeaux, LabEx COTE, Integrat & Theoret Ecol Grp, F-33615 Pessac, France
[5] Ctr Biodivers Theory & Modelling, Theoret & Expt Ecol Stn, CNRS, 2 Route CNRS, F-09200 Moulis, France
[6] Univ Canterbury, Sch Biol Sci, Bioprotect Ctr & Ctr Integrat Ecol, Private Bag 4800, Christchurch, New Zealand