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Sun Yat Sen Univ, Pearl River Estuary Marine Ecosyst Res Stn, Guangdong Prov Key Lab Marine Resources & Coastal, Zhuhai Key Lab Marine Bioresources & Environm,Sch, Zhuhai 519082, Peoples R China Sun Yat Sen Univ, Pearl River Estuary Marine Ecosyst Res Stn, Guangdong Prov Key Lab Marine Resources & Coastal, Zhuhai Key Lab Marine Bioresources & Environm,Sch, Zhuhai 519082, Peoples R China