a Department of Biology, Universit´e de Fribourg, Chemin du Mus´ee 10, CH-1700 Fribourg, Switzerland b Department of Mathematics, Universit´e de Fribourg, Chemin du Mus´ee 23, CH-1700 Fribourg, Switzerland c Swiss Institute of Bioinformatics, 1700 Fribourg, Switzerland d Aire de Conservation de Chinko, African Parks RCA,Bangui,中非共和国E工程与管理学院VAUD,HES-SO,应用科学与艺术大学西方瑞士