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abundances of P. ciliata had more diverse meiofaunal and macrofaunal communities compared with areas of low abundances, while Reise (1983b) found that the presence of dense assemblages of P. elegans promoted the abundance of small benthic organisms by approximately 40%. Morgan (1997), using a correlation approach, indicated that the majority of the most common taxa in the Baie de Somme, France, e.g., Eteone longa, Hediste diversicolor and Cerastoderma edule, were significantly positively correlated with P. elegans abundances. Many studies endeavouring to elucidate the mechanism by which high numbers of tube-builders affect infaunal community structure have focused on the way in which recolonisation is effected (e.g., Gallagher et al., 1983; Trueblood, 1991). Resident adults may influence colonists in many ways determined by their feeding mode and modifications to the sediments and hydrodynamics (Thrush et al., 1992). However, experimental studies on such mechanisms have proved equivocal. Since recruitment is usually assayed some time after settlement and metamorphosis (Bachelet, 1990) the actual patterns of larval settlement are often obscured (Hadfield, 1986). Consequently, studies have failed to assess the mechanisms by which tube-builders affect recolonisation since it is inherently difficult to distinguish between differential settlement and differential mortality of larvae (Woodin, 1986). Larvae of some marine benthic invertebrate species have been experimentally shown to actively select settlement sites using certain cues (Scheltema, 1974; Woodin, 1986; Butman et al., 1988a; Pawlik and Butman, 1993; Hsieh, 1994). However, it is likely that in the field, water flow is greater than the swimming speeds of larvae and instead, larvae are transported as passive particles and deposited via passive entrainment (Hannan, 1984; Butman, 1987; Butman et al., 1988b). Eckman (1983) predicted that velocity of near-bed flow through plastic straws (seagrass mimics) would be reduced to such an extent that suspended particles, such as larvae and meiofauna, would be passively deposited. His results supported this prediction although his experimental design was flawed by replication of only one of his treatments, his defaunated control. Later experiments have incorporated replication of the defaunated controls but the results have been inconclusive due to a number of factors including a plot size too small to allow the formation of a fully developed boundary layer (Kern and Taghon, 227

1986) and low level of replication resulting in too low a statistical power (Ragnarrson, 1996). The present study was observational and not intended to determine the mechanisms responsible for any differences in abundances between patch and non-patch communities. However, the increased silt/clay fraction and higher meiofaunal abundances in patches may have been due to increased passive deposition due to the reduction in velocity of near-bed flow. In the same way, passive larval entrainment may be responsible for some of the observed differences in this study. This was observed even though tube-building spionids have been shown to ingest settling bivalve larvae (Breese and Phibbs, 1972; Daro and Polk, 1973). Once an individual of a species had colonised a P. elegans patch, the physico-chemical effects of the tubes possibly concurred to provide an increased food supply in the form of the flourishing microbial and meiofaunal communities observed in this study. Furthermore, the resistance to shearing forces provided by the beds may have allowed a dense community by virtue of individuals not being 'swept' away (Morgan, 1997). Increased abundances of meiofauna (43%) were observed by Reise (1983b) in P. elegans patches compared to areas lacking the spionid, whilst a similar increase has been documented for beds of another spionid, Polydora ciliata, by Noji (1994), who suggested that the meiofauna were utilising the worms faecal pellets as a food source. The presence of C. volutator in patches, while almost completely absent outside patches, probably resulted from active habitat selection of adults to areas of increased silt/clay fraction and more stabilised sediments. This amphipod attains particularly high densities in muddy sediments with high numbers of diatoms (Lawrie, 1996). Its almost exclusive existence within patches suggests some sort of functional group interaction (sensu Woodin, 1976) for this tube-building, deposit-feeding amphipod. Similarly, Reise (1978) noticed C. volutator beds harbouring large numbers of P. elegans in the Wadden Sea. Sediment differences between P. elegans patches and non-patch areas. Sediment analyses indicated that although sediment water content was the same for patch and non-patch areas, the former consistently contained significantly increased 228

abundances of P. ciliata had more diverse meiofaunal and macrofaunal communities<br />

compared with areas of low abundances, while Reise (1983b) found that the presence<br />

of dense assemblages of P. elegans promoted the abundance of small benthic<br />

organisms by approximately 40%. Morgan (1997), using a correlation approach,<br />

indicated that the majority of the most common taxa in the Baie de Somme, France,<br />

e.g., Eteone longa, Hediste diversicolor and Cerastoderma edule, were significantly<br />

positively correlated with P. elegans abundances.<br />

Many studies endeavouring to elucidate the mechanism by which high numbers of<br />

tube-builders affect infaunal community structure have focused on the way in which<br />

recolonisation is effected (e.g., Gallagher et al., 1983; Trueblood, 1991). Resident<br />

adults may influence colonists in many ways determined by their feeding mode and<br />

modifications to the sediments and hydrodynamics (Thrush et al., 1992). However,<br />

experimental studies on such mechanisms have proved equivocal. Since recruitment<br />

is usually assayed some time after settlement and metamorphosis (Bachelet, 1990) the<br />

actual patterns of larval settlement are often obscured (Hadfield, 1986).<br />

Consequently, studies have failed to assess the mechanisms by which tube-builders<br />

affect recolonisation since it is inherently difficult to distinguish between differential<br />

settlement and differential mortality of larvae (Woodin, 1986).<br />

Larvae of some marine benthic invertebrate species have been experimentally shown<br />

to actively select settlement sites using certain cues (Scheltema, 1974; Woodin, 1986;<br />

Butman et al., 1988a; Pawlik and Butman, 1993; Hsieh, 1994). However, it is likely<br />

that in the field, water flow is greater than the swimming speeds of larvae and instead,<br />

larvae are transported as passive particles and deposited via passive entrainment<br />

(Hannan, 1984; Butman, 1987; Butman et al., 1988b). Eckman (1983) predicted that<br />

velocity of near-bed flow through plastic straws (seagrass mimics) would be reduced<br />

to such an extent that suspended particles, such as larvae and meiofauna, would be<br />

passively deposited. His results supported this prediction although his experimental<br />

design was flawed by replication of only one of his treatments, his defaunated control.<br />

Later experiments have incorporated replication of the defaunated controls but the<br />

results have been inconclusive due to a number of factors including a plot size too<br />

small to allow the formation of a fully developed boundary layer (Kern and Taghon,<br />

227

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