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The sole reliance on a planktonic mode of recruitment has implications for not only the smaller scale (metres) heterogeneity of this species on Drum Sands (Chapter 2) but also for the larger-scale maintenance of high population densities on Drum Sands in view of very high losses of planktonic larvae (Thorson, 1950; Mileikovsky, 1971; Vance, 1973; Bachelet, 1990). Morgan (1997) suggested that for the P. elegans population in the Baie De Somme, population maintenance was facilitated by the larval retentative properties of the bay. Active migration in the water column, which could enable a species to take advantage of directional water flows, has not been shown for polychaete larvae and therefore passive retentive mechanisms must be responsible for population maintenance in some areas (Morgan, 1997). The differences in passive larval retention between bays and the open coast was investigated by Gaines and Bertness (1992). They found that during periods of long flushing times, recruitment of barnacles within Naragansett Bay, Rhode Island, was significantly higher than recruitment to nearby coastal sites, while during periods of short flushing times, this difference was not observed. Tidally-induced retainment of spionid larvae have been shown in many bays and estuaries (see Morgan, 1997). Therefore, it is possible for passively dispersing larvae to be retained within their bay or estuary of origin by the hydrodynamic characteristics of the area and thus the possibility of population maintenance by a dispersive mode of reproduction is increased. There is evidence to suggest that the planktonic larvae of the P. elegans population on Drum Sands may be retained within the area to a certain extent. Dyke (1987) proposed that within the Firth of Forth, tidal residuals are small and that total residuals are more due to wind than the tide. Craig (1972) indicated that the residence time for seawater within the area was about 8 months. This suggests that the flushing time within the Firth of Forth is relatively long and, therefore, larval retention is likely to be high. Furthermore, Covill (1972, 1975) found that bays or indented coastlines along the southern Firth of Forth produced eddy currents which tended to distort the general tidal flow pattern of the Firth of Forth and that this effect was exacerbated by freshwater flows from rivers (Covill, 1972). The indented shape of Drum Sands, together with the added effect of the River Almond, suggests that eddy currents 79

probably exist around Drum Sands which may help retain P. elegans larvae. Furthermore, since the largest P. elegans populations within the Forth are found within the less saline Forth Estuary (SEPA, pers. comm.), planktonic larvae produced by them are likely to pass Drum Sands since this water passes along the south shore of the Forth (Dyke, 1987). This could be another potential input of larvae for the Drum Sands population. However, the mode of reproduction of P. elegans within the less saline Forth Estuary has not been studied. The importance of pelagic larvae for shaping community structure has been discussed by Lewin (1986). He suggested that previous models of marine benthic community structuring, sensu Paine and Connell, have overlooked the importance of the larval fluxes since these models were based on experiments carried out where larval influxes were saturating. In areas where larval settlement is saturating, evidence suggests that communities are shaped by post-settlement processes such as predation and competition. In areas where larval recruitment is not saturated however, larval-supply fluctuations become more important in shaping demographic processes (Lewin, 1986; Bachelet, 1990). Therefore, it is possible that the P. elegans population at Drum Sands is structured by fluxes in larval settlement and the processes which effect it, rather than processes acting on the adult population. Consequently, the effect of larval recruitment on population density changes is specifically addressed in this study in response to a number processes including macroalgal mat establishment (Chapters 4 and 5), small-scale disturbances (Chapter 6) and the generation of micro-scale spatial heterogeneity due to adult-juvenile interactions (Chapter 7). However, further studies have to be carried out in order to assess whether P. elegans larval recruitment is indeed saturated or not on this sandflat. Since the life history of a species governs that species' dispersal dynamics and recruitment in a particular environment, Levin (1984a) suggested that it determined the scale of disturbance a species was potentially capable of exploiting. She noted that polychaetes with small adult size, brood protection, small brood size and reduced larval phases (e.g., Capitella spp., Streblospio benedicti) were adapted to small-scale disturbances such as those caused by ray foraging and human pit-digging, while polychaetes with larger brood sizes and longer-lived larvae, whose planktonic 80

The sole reliance on a planktonic mode of recruitment has implications for not only<br />

the smaller scale (metres) heterogeneity of this species on Drum Sands (Chapter 2) but<br />

also for the larger-scale maintenance of high population densities on Drum Sands in<br />

view of very high losses of planktonic larvae (Thorson, 1950; Mileikovsky, 1971;<br />

Vance, 1973; Bachelet, 1990). Morgan (1997) suggested that for the P. elegans<br />

population in the Baie De Somme, population maintenance was facilitated by the<br />

larval retentative properties of the bay. Active migration in the water column, which<br />

could enable a species to take advantage of directional water flows, has not been<br />

shown for polychaete larvae and therefore passive retentive mechanisms must be<br />

responsible for population maintenance in some areas (Morgan, 1997). The<br />

differences in passive larval retention between bays and the open coast was<br />

investigated by Gaines and Bertness (1992). They found that during periods of long<br />

flushing times, recruitment of barnacles within Naragansett Bay, Rhode Island, was<br />

significantly higher than recruitment to nearby coastal sites, while during periods of<br />

short flushing times, this difference was not observed. Tidally-induced retainment of<br />

spionid larvae have been shown in many bays and estuaries (see Morgan, 1997).<br />

Therefore, it is possible for passively dispersing larvae to be retained within their bay<br />

or estuary of origin by the hydrodynamic characteristics of the area and thus the<br />

possibility of population maintenance by a dispersive mode of reproduction is<br />

increased.<br />

There is evidence to suggest that the planktonic larvae of the P. elegans population on<br />

Drum Sands may be retained within the area to a certain extent. Dyke (1987)<br />

proposed that within the Firth of Forth, tidal residuals are small and that total residuals<br />

are more due to wind than the tide. Craig (1972) indicated that the residence time for<br />

seawater within the area was about 8 months. This suggests that the flushing time<br />

within the Firth of Forth is relatively long and, therefore, larval retention is likely to be<br />

high. Furthermore, Covill (1972, 1975) found that bays or indented coastlines along<br />

the southern Firth of Forth produced eddy currents which tended to distort the general<br />

tidal flow pattern of the Firth of Forth and that this effect was exacerbated by<br />

freshwater flows from rivers (Covill, 1972). The indented shape of Drum Sands,<br />

together with the added effect of the River Almond, suggests that eddy currents<br />

79

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