[1] Univ Sheffield, Dept Anim & Plant Sci, Sheffield S10 2TN, S Yorkshire, England
[2] Univ Leeds, Sch Biol, Fac Biol Sci, Leeds LS2 9JT, W Yorkshire, England
[3] Univ Groningen, Groningen Inst Evolutionary Life Sci, Behav Ecol & Physiol Grp, POB 11103, NL-9700 CC Groningen, Netherlands
[4] Univ East Anglia, Sch Biol Sci, Norwich Res Pk, Norwich NR4 7TJ, Norfolk, England
[5] Nat Seychelles, POB 1310, Mahe, Seychelles