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of tubes on near-bed flow and the indirect effects of the resulting biogenic changes<br />

(Lukenbach, 1986).<br />

Polychaete tubes are thought to exert important and complex effects on near-bed flow.<br />

While isolated tubes, or 'roughness elements' in hydrodynamic terms (Eckman, 1983),<br />

destabilise sediments by creating local currents which result in sediment scour and<br />

local deposition (Eckman et al., 1981; Lukenbach, 1986; 1987), there is a widely<br />

accepted notion that sediments are stabilised by arrays of tubes. This opinion has<br />

developed from both field studies (Sanders et al., 1962; Fager, 1964; Daro and Polk,<br />

1973) and laboratory flume studies (Nowell and Church, 1979; Nowell et al., 1981;<br />

Eckman, 1983). Tube size, geometry, and numerical density determine the properties<br />

of near-bed flow, including the magnitude of the shear stress exerted on the bed (and<br />

hence its susceptibility to erosion, i.e., stability), the rate of fluid transport, and the<br />

production of turbulence (Eckman et al., 1981; Nowell et al., 1981). The shear stress<br />

exerted on the bed affects emigration of established individuals and the composition<br />

of the sediments, while fluid transport near the bed will affect immigration rates of<br />

fauna dispersed passively by lateral advection. It is conceivable that these<br />

hydrodynamic processes alone may result in the distinct faunal communities observed<br />

in field studies. The effects of tube-beds on fluid transport have been shown to<br />

facilitate colonisation by micro-organisms which further helps stabilise the sediments<br />

(Eckman et al., 1981; Lukenbach, 1986) by mucus binding (Grant et al., 1986). Since<br />

the velocity of near-bed flow generally exceeds the swimming speeds of larvae (Chia,<br />

1989), larvae are generally advected horizontally as passive particles along the bed<br />

(Butman, 1987) and deposition to the bottom is usually predominantly via passive<br />

entrainment (Hannan, 1984; Pawlik and Butman, 1993; Snelgrove et al., 1993).<br />

Eckman (1983) however, using flume experiments suggested that the velocity of near-<br />

bed flow was reduced using seagrass mimics to such an extent that larvae would be<br />

passively deposited. In addition to the hydrodynamic effect of their tubes, tube-<br />

building organisms themselves significantly affect processes at the sediment-water<br />

interface by their behaviour, e.g., feeding, burrowing, defecation and tube irrigation<br />

(Rhoads and Young, 1970; Frithsen and Doering, 1986; Noji and Noji, 1991).<br />

202

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