[1] Univ Leeds, Fac Biol Sci, Sch Biol, Leeds, W Yorkshire, England
[2] Univ Sheffield, Dept Anim & Plant Sci, NERC Environm Omics Visitor Facil, Sheffield, S Yorkshire, England
[3] Univ Groningen, Groningen Inst Evolutionary Life Sci, Groningen, Netherlands
[4] Univ Oxford, Recanati Kaplan Ctr, Dept Zool, Wildlife Conservat Res Unit, Abingdon, Oxon, England
[5] Univ British Columbia, Dept Biol, Kelowna, BC, Canada