[1] Carnegie Inst Sci, Dept Global Ecol, 260 Panama St, Stanford, CA 94305 USA
[2] Sci Serv, South African Natl Pk,Private Bag X402, ZA-1350 Skukuza, South Africa
[3] Univ Witwatersrand, Sch Anim Plant & Environm Sci, Ctr African Ecol, Private Bag 3, ZA-2050 Johannesburg, South Africa
[4] Max Planck Inst Biogeochem, Dept Biogeochem Proc, Hans Knoll Str 10, DE-07745 Jena, Germany