[1] Univ Helsinki, Fac Biol & Environm Sci, Doctoral Programme Wildlife Biol, Viikinkaari 1,POB 65, Helsinki 00014, Finland
[2] Univ Helsinki, Fac Biol & Environm Sci, Res Programme Organismal & Evolutionary Biol, Viikinkaari 1,POB 65, Helsinki 00014, Finland
[3] Univ Helsinki, HiLIFE Helsinki Inst Life Sci, Viikinkaari 1,POB 65, Helsinki 00014, Finland
[4] Macquarie Univ, Sch Nat Sci, 14 Eastern Rd, Sydney, NSW 2109, Australia