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Other theories have been postulated for the observed community differences within tube-beds. Woodin (1978, 1981) suggested that the observed effects result from the 'refuge' provided by the high numbers of tubes. Both predation and local disturbances have been shown to be important structuring forces for soft-bottom infauna (Reise, 1978; Woodin, 1978; Zajac and Whitlatch, 1982a; Ambrose, 1984; Thrush, 1988; Hall et al., 1993) and Woodin (1978) suggested that refuges may be created by organisms whose structures buffer the impact of physiological stress or inhibit the access of predators. Woodin (1976) viewed soft-bottom community structuring from the perspective of 'functional group' interactions in which 'tube- dweller', `bioturbator' and `suspension-feeder' functional groups should each be unfavourable to each other. This 'functional group' interaction hypothesis implied that the maintenance of discrete dense assemblages in infaunal systems was due to interactions between established infaunal individuals and settling larvae. Therefore, a tube-building species would be expected to exhibit increased densities within an assemblage of other tube-builders by this interaction, whilst bioturbators and suspension-feeders would be expected to exhibit lower densities. The main tube-building polychaete species forming dense tube-beds in northern European shallow waters tend to be either Lanice conchilega or Pygospio elegans. Several studies such as Eagle (1973, 1975), Carey (1982) and Ragnarsson (1996) have described the fauna within L. conchilega tube-beds, however, relatively few studies have focused on the beds formed by P. elegans (e.g., Dupont, 1975; Morgan, 1997). The dense tube-beds at Drum Sands allowed an investigation of the effects of this tube-builder on an intertidal sandflat. Specifically, the differences in the biology and the sediments within P. elegans patches and non-patch areas of sand were studied. 203

METHODS The sampling for the investigation described in Chapter 6 was carried out at the same time and within the same patch and non-patch plots as those for the present study. However, a further survey was also carried out in August 1998 for this study. Faunal sampling methods, sample treatment and sorting, faunal identification and statistical analyses for this chapter are as described in Chapter 6. While sediment variables were only measured on one occasion for the investigation described in Chapter 6, the same variables were measured at each sample date for the present study, i.e., April, August and December 1997, and August 1998. In addition to the size measurements of P. elegans, individuals of other species with mean abundances above 20 per core in patch and non-patch samples were measured. P. elegans and C. capitata were measured as 5th setiger and 3rd setiger widths respectively. Small bivalves were measured as shell width using a microscope (x20 magnification) fitted with an eyepiece graticule while larger ones were measured with a pair of calipers. 204

Other theories have been postulated for the observed community differences within<br />

tube-beds. Woodin (1978, 1981) suggested that the observed effects result from the<br />

'refuge' provided by the high numbers of tubes. Both predation and local<br />

disturbances have been shown to be important structuring forces for soft-bottom<br />

infauna (Reise, 1978; Woodin, 1978; Zajac and Whitlatch, 1982a; Ambrose, 1984;<br />

Thrush, 1988; Hall et al., 1993) and Woodin (1978) suggested that refuges may be<br />

created by organisms whose structures buffer the impact of physiological stress or<br />

inhibit the access of predators. Woodin (1976) viewed soft-bottom community<br />

structuring from the perspective of 'functional group' interactions in which 'tube-<br />

dweller', `bioturbator' and `suspension-feeder' functional groups should each be<br />

unfavourable to each other. This 'functional group' interaction hypothesis implied<br />

that the maintenance of discrete dense assemblages in infaunal systems was due to<br />

interactions between established infaunal individuals and settling larvae. Therefore, a<br />

tube-building species would be expected to exhibit increased densities within an<br />

assemblage of other tube-builders by this interaction, whilst bioturbators and<br />

suspension-feeders would be expected to exhibit lower densities.<br />

The main tube-building polychaete species forming dense tube-beds in northern<br />

European shallow waters tend to be either Lanice conchilega or Pygospio elegans.<br />

Several studies such as Eagle (1973, 1975), Carey (1982) and Ragnarsson (1996) have<br />

described the fauna within L. conchilega tube-beds, however, relatively few studies<br />

have focused on the beds formed by P. elegans (e.g., Dupont, 1975; Morgan, 1997).<br />

The dense tube-beds at Drum Sands allowed an investigation of the effects of this<br />

tube-builder on an intertidal sandflat. Specifically, the differences in the biology and<br />

the sediments within P. elegans patches and non-patch areas of sand were studied.<br />

203

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