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likely that the tube structure of dead worms remain intact for some time after the<br />

occupant has died. In this way, the direct hydrodynamic effect of P. elegans tubes<br />

within patches in this study may have been far greater than that estimated from worm<br />

densities alone.<br />

Sediment stabilisation by P. elegans has been previously demonstrated by shear<br />

strength measurements (Meadows and Hariri, 1991; Morgan, 1997). Although such<br />

measurements were not taken in this study there is strong evidence to suggest that the<br />

sediments within P. elegans patches on Drum Sands were stabilised, i.e., the lack of<br />

ripple marks in an otherwise rippled area, the increased silt/clay fraction, and the<br />

presence of a golden hue, assumed to be diatoms, on the sediment surface. Eckman et<br />

al. (1981) suggested that these observations, especially the latter, provided strong<br />

evidence of sediment stabilisation. In the studies by Sanders et al. (1962) and Fager<br />

(1964), sediment stabilisation was also assumed by the observations of a golden hue,<br />

confirmed to be diatoms by chlorophyll analysis in the former study, and the absence<br />

of ripple marks where highest tube densities were found. The polychaete densities<br />

producing these effects in their studies were also far below those expected for<br />

sediment stabilisation using the empirical formula of Nowell and Church (1979), i.e.,<br />

observed densities being 600 C. torquatalm2 and 500-1,000 0. fusiformislm2 in the<br />

studies by Sanders eta!. (1962) and Fager (1964), respectively. For example, Eckman<br />

et al. (1981) calculated that a density of at least 14,500 0. fusiformislm2 was<br />

necessary for sediment stabilisation.<br />

A number of studies have experimentally shown that extracellular mucus films<br />

produced by macrofauna during tube construction (Meadows and Tufail, 1986), and<br />

diatoms can stabilise sediments (Frostick and McCave, 1979; Grant et al., 1986;<br />

Tufail eta!., 1989; Paterson, 1994). Sediments around tubes have been shown to have<br />

increased numbers of diatoms (Sanders et al., 1962) and meiofauna (Eckman, 1983;<br />

Noji, 1994). The dense films of diatoms observed on the P. elegans patch surfaces<br />

during this study support the contention that indirect sediment stabilisation was<br />

possibly appreciable. Therefore, the observed differences in colonisation between<br />

patches and non-patches in this study probably resulted from differences in sediment<br />

stability.<br />

166

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