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Vertical zonation and aggregated distribution of the Manila clam on subtidal sand flats in a coastal brackish lagoon along the Sea of Japan

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Abstract

The Manila clam, Ruditapes philippinarum, has maintained small-sized populations in a semi-enclosed brackish lake along the Sea of Japan, the Honjo area of Lake Nakaumi, although the environment and biota of this area have changed dramatically due to a large-scale reclamation project. There should be underlying processes that enable the restoration of this species from small-sized populations, such as the existence of source (i.e. reproductive) populations in other areas and depth zones of the lake. However, there has been no robust, properly designed evaluation of the distribution of the Manila clam in the subtidal sand flats. In order to elucidate the possible mechanisms that allow for the persistence of populations of the Manila clam, we examined the spatiotemporal and vertical variation in distributions of 0-age clams in the subtidal zone of sand flats. Seasonal effects on population variations showed erratic changes among depth zones without a decreasing trend along the depth gradient. Further, many local populations became extinct even in the shallower zones due to seasonal (summer) hypoxia at deeper zones and hypoxia by the accumulation of key benthic species (Asian mussel and decaying macroalgae) in mats at shallower zones. A few surviving local populations were stable with a spatial-fragmental (patchy) distribution, associated with fragmented accumulations of Asian mussels and macroalgae. Efforts to maintain stable populations and to restore this species in the subtidal area may depend on these small, restricted, patchy local populations. These findings suggest that high fertility and productivity of the Manila clam as well as patchy distribution of small populations may contribute to the maintenance of the population and the avoidance of extinction (by spatially diffusing the risk of extinction) in harsh environments resulting from the reclamation project.

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