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touched by M. balthica siphons. C. edule has been shown to inhale settling P. elegans<br />

larvae (Noyer, 1993; cited by Morgan, 1997). Reise (1983c; 1985), Jensen (1985) and<br />

Flach (1996) have suggested that C. edule could affect other infaunal species by<br />

disturbing the upper sediment layer due to its crawling (ploughing) and 'shaking'<br />

(Flach, 1996) behaviour. In Flach's study, significant reductions in densities of most<br />

infaunal species, including P. elegans, were found even at the lowest C. edule density,<br />

i.e., 4-8% C. edule occupancy (area occupied by C. edule). The C. edule occupancy in<br />

P. elegans patches in August 1998 was calculated to be 9.5%, with an appreciably<br />

greater 'disturbance area' (i.e., total area disturbed) (Flach, 1996). Therefore, it is<br />

likely that C. edule was having a large negative effect on P. elegans within patches in<br />

August 1998 due to sediment disturbance.<br />

Settling spionid larvae seem to be particularly prone to the disturbance and predation<br />

effects of C. edule and in most studies it is the juveniles which have suffered the<br />

greatest decrease in numbers. Size measurements taken of P. elegans from the August<br />

1998 samples in this study showed that although the size-frequency distributions were<br />

not quite significantly different, the proportion of juveniles was greatly reduced in P.<br />

elegans patches compared with non-patches which had significantly lower bivalve<br />

populations. Therefore, the decrease in numbers of P. elegans in patches at this time<br />

was mainly due to the juveniles, suggesting that there was high larval mortality in<br />

patches during the peak settlement period earlier in the year. However, it is not<br />

possible to determine whether this was predominantly due to sediment disturbance or<br />

inhalation by the bivalves.<br />

It could be envisaged that the numerical dominance of C. edule, and to a lesser extent<br />

M. balthica, were likely to cause further reductions in P. elegans densities from those<br />

of August 1998 because of their negative effects on spionids. The C. edule<br />

individuals sampled in August 1998 had reached a mean size of 2.5mm length and<br />

hence had already reached a size refuge from predation by infaunal predators such as<br />

Nereis spp. and epibenthic species such as Crangon crangon, 0-group Carcinus<br />

maenas and 0- and I-group Pleuronectes spp. (Reise, 1985). C. maenas and C.<br />

crangon have been shown to be important predators on juvenile C. edule, and C.<br />

crangon, found in moderate abundances during sampling at Drum Sands, has been<br />

232

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