[1] Univ Cadiz, Dept Biol, IVAGRO, Campus Rio San Pedro, Puerto Real 11510, Spain
[2] Univ Zurich, Dept Evolutionary Biol & Environm Studies, Winterthurerstr 190, CH-8057 Zurich, Switzerland
[3] Univ Sheffield, Dept Anim & Plant Sci, Alfred Denny Bldg,Western Bank, Sheffield S10 2TN, S Yorkshire, England
[4] Univ Queensland, Sch Biol Sci, Goddard Bldg 8, St Lucia, Qld 4072, Australia
[5] Max Planck Inst Demog Res, Evolutionary Demog Lab, Konrad Zuse Str 1, D-18057 Rostock, Germany