Relative importance of predation and intraspecific competition in regulating growth and survival of juveniles of the soft-shell clam, Mya arenaria L., at several spatial scales

Predation appears to be the single most important biotic factor regulating populations of bivalves in estuarine and coastal soft sediments. However, the relative roles of predation and intraspecific competition are rarely investigated simultaneously over different spatial scales, making generalities about these mechanisms difficult. Using juveniles of the soft-shell clam, Mya arenaria (initial mean shell length [SL] +/- 95% CI = 12.4 +/- 0.13 mm), I tested the interactive effects of predator deterrence and intraspecific density (660 vs. 1320 individuals m(-2)) on growth and survival responses over a 185-day period from May to November 2003 at spatial scales that spanned four orders of magnitude: embayments, sites within embayments, tidal gradients, and blocks that were 10,000's, 1000's, 100's, and 5 m apart, respectively. Replicate field experiments were conducted from May to November 2003 at the upper and lower tidal heights at each of two intertidal mud flats (sites) within each of two embayments (Passamaquoddy Bay [PB] and Cobscook Bay [CB]) in eastern Maine. Mean survival, relative growth, and the abundance of wild recruits each varied significantly over all spatial scales. Predation was the most important factor affecting clam survival, explaining 45% of the total variability, whereas embayment, sites within embayments, tidal gradient, and intraspecific density collectively accounted for less than 10% of the variation. At all four intertidal sites, clam survival in experimental units designed to deter predators averaged 72%, but the degree of enhancement varied between embayments (PB = 6 1 %; CB = 267%). Average survival rate was higher (by 12-16%), but growth was slower (by ca. 50%) in upper vs. lower intertidal blocks; however, the patterns differed for both variables between sites within each embayment. The effect of increasing intraspecific clam density was to lower survival by ca. 17% (56% [660 m(-2)] vs. 48% [1320 m(-2)]) in both embayments, but growth was unaffected. Overall, clams doubled in SL, although mean relative growth was 15% greater in CB than PB. Tidal gradient, sites within embayments, and blocks were the three most important factors explaining 35%, 19%, and 22% of total variation in relative clam growth, respectively. In Maine and the northeast US, juveniles of Mya reach their highest abundance above mean low tide levels. Experimental evidence presented here suggests that differential predation along the tidal gradient is the dominant factor controlling clam abundance and distribution patterns in the intertidal zone. (c) 2006 Elsevier B.V. All rights reserved.