[1] Australian Natl Univ, Res Sch Earth Sci, Canberra, ACT 0200, Australia
[2] Australian Natl Univ, Res Sch Biol Sci, Canberra, ACT 0200, Australia
[3] Harry Oppenheimer Okavango Res Ctr, Maun, Botswana
[4] Max Planck Inst Biogeochem, D-07701 Jena, Germany