[1] Bournemouth Univ, Fern Barrow, Talbot Campus, Poole BH12 5BB, Dorset, England
[2] Univ Paris 06, MNHN, IRD, UMR BOREA, F-75231 Paris 5, France
[3] UMMISCO, IRD UPMC 209, Ctr Math & Computat Modeling Complex Syst, Int Res Unit, F-93143 Bondy, France