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In April, when P. elegans larval availability in the water column was at its highest on Drum Sands, significantly higher numbers of P. elegans larvae colonised azoic sediments in patches compared with non-patch azoic sediments. Many laboratory experimental studies have been carried out to elucidate the precise mechanisms by which differential larval settlement of infaunal species is achieved (Scheltema, 1974; Woodin, 1986; Butman et al., 1988a; Pawlik and Butman, 1993; Hsieh, 1994) but such studies on polychaetes have mainly concentrated on families other than Spionidae, e.g., Capitellidae (Butman et al., 1988b; Grassle and Butman, 1989) and Sabellidae (e.g., Pawlik et al., 1991) and it is likely that mechanisms are species- specific. However, settlement cues associated with adult populations were unlikely to have been responsible for the observed differences in P. elegans larval recruitment in the present study because colonisation occurred on totally defaunated sediments. It is possible that larval behavioural responses to local boundary-layer flow may have produced the observed differences for juvenile P. elegans in this study. Such behaviour has been demonstrated for the sabellid Phragmatopoma lapidosa califomica (Pawlik et al., 1991) and it was proposed that this enabled this species to form large tube-beds. Savidge and Taghon (1988) showed that in the field, colonisation was dominated by passive advection of larvae (see review by Olafsson et al., 1994). Eckman (1983) suggested that passive larval entrainment can be higher in areas of stabilised sediments. Increased larval entrainment was suggested to have been a possible reason why C. edule spat were present in higher numbers in P. elegans patches on Drum Sands (Chapter 8). Increased passive entrainment to P. elegans patches could, therefore, have been responsible for the increased recruitment of P. elegans larvae to patches in this study. However, one must be cautious, apparent increased recruitment to patches could equally have resulted from increased erosion of settled larvae in non- patch areas. Since samples were taken some time after settlement, inferring settlement patterns from observed recruitment patterns can be potentially misleading (Hadfield, 1986; Woodin, 1986). Woodin (1986) suggested that to differentiate between larval settlement and recruitment in field studies is inherently very difficult. Differential 167

mortality and/or erosion were likely to have been factors in this study, i.e., those larvae settling into patches are less likely to have been eroded in the stabilised environment of patches and presumably benefit from the increased food supply. Zajac and Whitlatch (1982a) demonstrated that initial colonisation following small- scale sediment disturbances were predominantly dependent on the temporal changes in ambient populations. Although such temporal changes in the ambient communities on Drum Sands were important in initial colonisation and presumably in later successional dynamics in the present study, the interspersion of the two plot types demonstrated that the early stages of succession can vary at small spatial scales concurrently. Does the successional stage of a community affect the colonisation mode of early colonisers? Two species, i.e., the polychaetes P. elegans and C. capitata, colonised in numbers sufficiently high for size-frequency analyses. Although no significant differences were observed in the size distributions of P. elegans colonising azoic sediments of the two habitat types during April 1997, significant differences during August and December 1997 were observed due to higher numbers of adults colonising patch azoic sediments relative to non-patch sediments. There were significant differences between the size distributions of C. capitata colonising the two habitat types in December 1997 which suggested that colonisation in patch azoics was predominantly via adults while that to non-patch azoics was via larvae. The results of the present experiments, therefore, indicated that in addition to affecting initial community composition following a disturbance, dense assemblages of P. elegans also affected the colonisation mode of some species. These patches of increased P. elegans densities reflected different successional stages from surrounding sediments (Noji and Noji, 1991). Many studies have found that post-larval immigration can play a substantial role in colonisation after a small-scale disturbance (Dauer and Simon, 1976a; Levin, 1984a; Frid, 1989; Smith and Brumsickle, 1989; Wilson, 1992; 1994). This has been found 168

In April, when P. elegans larval availability in the water column was at its highest on<br />

Drum Sands, significantly higher numbers of P. elegans larvae colonised azoic<br />

sediments in patches compared with non-patch azoic sediments. Many laboratory<br />

experimental studies have been carried out to elucidate the precise mechanisms by<br />

which differential larval settlement of infaunal species is achieved (Scheltema, 1974;<br />

Woodin, 1986; Butman et al., 1988a; Pawlik and Butman, 1993; Hsieh, 1994) but<br />

such studies on polychaetes have mainly concentrated on families other than<br />

Spionidae, e.g., Capitellidae (Butman et al., 1988b; Grassle and Butman, 1989) and<br />

Sabellidae (e.g., Pawlik et al., 1991) and it is likely that mechanisms are species-<br />

specific. However, settlement cues associated with adult populations were unlikely to<br />

have been responsible for the observed differences in P. elegans larval recruitment in<br />

the present study because colonisation occurred on totally defaunated sediments.<br />

It is possible that larval behavioural responses to local boundary-layer flow may have<br />

produced the observed differences for juvenile P. elegans in this study. Such<br />

behaviour has been demonstrated for the sabellid Phragmatopoma lapidosa<br />

califomica (Pawlik et al., 1991) and it was proposed that this enabled this species to<br />

form large tube-beds.<br />

Savidge and Taghon (1988) showed that in the field, colonisation was dominated by<br />

passive advection of larvae (see review by Olafsson et al., 1994). Eckman (1983)<br />

suggested that passive larval entrainment can be higher in areas of stabilised<br />

sediments. Increased larval entrainment was suggested to have been a possible reason<br />

why C. edule spat were present in higher numbers in P. elegans patches on Drum<br />

Sands (Chapter 8). Increased passive entrainment to P. elegans patches could,<br />

therefore, have been responsible for the increased recruitment of P. elegans larvae to<br />

patches in this study. However, one must be cautious, apparent increased recruitment<br />

to patches could equally have resulted from increased erosion of settled larvae in non-<br />

patch areas. Since samples were taken some time after settlement, inferring settlement<br />

patterns from observed recruitment patterns can be potentially misleading (Hadfield,<br />

1986; Woodin, 1986). Woodin (1986) suggested that to differentiate between larval<br />

settlement and recruitment in field studies is inherently very difficult. Differential<br />

167

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