Spatial partitioning between species of the phytoplankton-feeding guild on an estuarine intertidal sand flat and its implication on habitat carrying capacity

The fishery yield of Manila clams, Ruditapes philippinarum, increased considerably in the 1970s but has decreased rapidly since the middle 1980s on extensive intertidal sand flats in Ariake Sound (Kyushu, Japan). A survey conducted in 2004 on a 3.4-km² sand flat located in the central part of the Sound (Shirakawa sand flat) revealed four dominant species: two thalassinidean shrimps (Upogebia major and Nihonotrypaea japonica), which are deep-reaching burrow dwellers with strong bioturbation activities, and two bivalves (Mactra veneriformis and R. philippinarum). All four species belong to a phytoplankton (diatom)-feeding guild. In the late 1970s, the Manila clam population prevailed in high densities over the entire sand flat, whereas its distribution was restricted to the lowest quarter of the shore in 2004. In contrast, the population sizes and zones of occurrence of the other phytoplankton feeders have expanded in the absence of R. philippinarum, perhaps an indication of competitive release. After establishment, effects of the thalassinidean shrimps on sediment stability appear to have further reduced clam abundances. Across the sand flat in 2004, wet weight population biomass estimates for N. japonica, U. major, M. veneriformis, and R. philippinarum (whole body for shrimps and soft tissue for bivalves) were 304, 111, 378, and 234 tonnes, respectively. Based on Manila clam fishery yield records from Shirakawa, the carrying capacity of the Shirakawa sand flat in the late 1970s was estimated to be two times greater than the sum value for the whole phytoplankton-feeding guild in 2004. It is hypothesized that (1) the amount of phytoplankton determines the carrying capacity for the benthic community on the Shirakawa sand flat, with both phytoplankton and benthic biomass at maxima in the late 1970s, and (2) the subsequent increases in competition for space have caused further declines in the Manila clam population biomass to approximately one-eighth of its past value.