[1] Minzu Univ China, Coll Life & Environm Sci, Beijing 100081, Peoples R China
[2] Chinese Acad Sci, Inst Bot, State Key Lab Vegetat & Environm Change, Beijing 100093, Peoples R China
[3] Chinese Acad Sci, Inst Geog Sci & Nat Resources Res, Lhasa Plateau Ecosyst Res Stn, Key Lab Ecosyst Network Observat & Modeling, Beijing 100101, Peoples R China
[4] San Diego State Univ, Biol Dept, San Diego, CA 92182 USA