Key Laboratory of Tropical Marine Bio-resources and Ecology (LMB), South China Sea Institute of Oceanology, Chinese Academy of Sciences, Guangzhou 510301, China; University of Chinese Academy of Sciences, Beijing 100049, China; Guangdong Provincial Key Laboratory of Applied Marine Biology (LAMB), Guangzhou, China; Southern Marine Science and Engineering Guangdong Laboratory (Guangzhou), China; Innovation Academy of South China Sea Ecology and Environmental Engineering, Chinese Academy of Sciences, Guangzhou 510301, China. Electronic address: tanyh@scsio.ac.cn.