[1] Univ Groningen, Groningen Inst Evolutionary Life Sci, Groningen, Netherlands
[2] Univ Lausanne, Dept Ecol & Evolut, Lausanne, Switzerland
[3] Mawana Game Reserve, Inkawu Vervet Project, Kwa Zulu, South Africa
[4] Wageningen Univ & Res, Biometris, Wageningen, Netherlands