The study area is located in the Jiulong River Estuary Mangrove Reserve
(117°54′E, 24°23′N) in the monsoonal subtropics of Southeast
China (Fig. 1). The climate of this region is warm and wet,with an average
annual temperature of 21 °C and average annual precipitation of
1371 mm, most of which falls between April and September. Tide pattern
in this region is typically semi-diurnal tides with an average tide
range of 2.98 m and seawater salinity of 17.12 ppt. The four vegetation
types, namely, native Kandelia forest, exotic Sonneratia forest, native
Cyperus marsh and exotic Spartina marsh are adjacent under the same
intertidal elevation and thus experience similar tidal influence (Fig. 1).