1. Department of Evolution, Ecology and Organismal Biology The Ohio State University Columbus OH 43210 USA
2. Smithsonian Tropical Research Institute Box 0843‐03092 Balboa Ancón Republic of Panama
3. Mathematical Biosciences Institute The Ohio State University Columbus OH 43210 USA
4. State Key Laboratory of Vegetation and Environmental Change Institute of Botany Chinese Academy of Sciences 20 Nanxincun Xiangshan Beijing 100093 China