Marsh periwinkles of 5 to 7 mm in shell height were eaten regularly by blue crabs. Fractures marking unsuccessful crab attacks were present in about 25% of medium-sized (10-16 mm) snails and over 60% of larger snails (over 16 mm). Medium-sized snails, subject to predation, leave the water more frequently than larger snails, but only about a quarter of the snail population leaves the water during high tide. We found no evidence that the snails leave the water because they sense blue crabs in the water. INTRODUCTION The marsh periwinkle littorina ivorata (Say) is a grayish gibbous, intertidal snail common in salt marshes from New York to Texas. It is believed to be a grazer, feeding largely on plant detritus and epipelic algae (Odum and Smalley 1959; Stiven and Kuenzler 1979). According to Bingham (1972) and Hamilton (1976, 1978), periwinkles are effectively supratidal; they avoid submergence by crawling upward out of the water when innundated. Bingham (1972) attributed the absence of snails in barren areas to “reluctance of the species to remain submerged.” Bleil and Gunn (1978) demonstrated that this response is not caused by threat of drowning; the snails can survive long periods under water. Hamilton (1976) attributed the avoidance of submergence to predation by the blue crab Callinectes sapidus Rathbun, which enters the marsh with the flood tide and preys on periwinkles. Crist (1979) found that a size gradient exists in periwinkles; larger individuals occur more frequently than smaller snails at lower elevations in the marsh. He attributed this in part to differential predation on smaller snails by blue crabs, which are more abundant at lower elevations. Preliminary observations on a study site at Wallops Island, Virginia, indicated that a substantial majority of specimens of L. irrorata remained under water at high tide, despite an abundance of emergent vegetation. That observation was an obvious contradiction to the behavior of L. irrorata in an aquarium-all snails crawl out of the water immediately (Bingham 1972). We noticed also that smaller snails tended to be more abundant out of water on stalks of vegetation than in the water. We reasoned that there may be a size-specific difference in the response of snails to submergence because of differential predation on smaller snails relative to larger ones. The purpose of this paper is to quantify the size-specific ‘Address reprint requests to Dr. Banta. Manuscript received February 8,1982; accepted July 28,1982. proportions of snails which emerge from the water at high tide in the field, and to test the hypothesis that blue crab predation may influence those proportions. MATERIALS AND METHODS The study site was a juvenile sloping foreshore marsh located on Cow Gut Flat at the north end of Wallops Island, near Chincoteague, Virginia. Drainage at low tide is nearly complete, and freshwater input is limited to rainfall and groundwater discharge. No tidal creeks or primary pans are present. Tall and medium vigor Spartina alterniflora (Loisel) predominate. Salicomia spp. is abundant at some higher elevations; the highest elevations are dominated by marsh elder, Iva fmtescens (Linnaeus) (Reidenbaugh and Banta 1980). Tidal wrack, consisting primarily of dead stalks of S. altemiflora, formed dense mats, which were rafted into the site during abnormally high tides (Reidenbaugh and Banta 1980). Vegetation compressed beneath stranded mats often is partly or completely killed, and in the most severe cases, secondary bare areas formed. Field experiments were conducted within a 17,000-m2 study site marked by wooden stakes placed at 10-m intervals from below mean low water to above mean high water. The study site has been named the Intensive Biometric Intertidal Survey (IBIS) marsh (Reidenbaugh and Banta 1980). To determine the size distribution of Littorina irrorata in the IBIS marsh survey grid, 30 sampling points were located relative to stakes, using computer-generated random numbers. Elevations at each collecting site were determined by linear interpolation among the four surrounding stakes. Stake elevations were determined by transit relative to Bench Mark IBIS, t 1.427 m National Geodetic Vertical Datum (NGVD). A 1-m2 quadrat was centered at each of the sampling points at or near the time of low tide between 16 and 18 July, 1979. The weather during this time was uniformly sunny and warm, with temperature maxima near 30°C. All marsh periwinkles were collected, counted, measured to the nearest 1 mm shell height, and returned to the same quadrat.
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