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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
- Page 44 and 45: 2.5 1.5 0.5 0 3 T (i) % Silt/clay%
- Page 46 and 47: v : m pattern Id pattern Ip pattern
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- Page 50 and 51: The results show that at the smalle
- Page 52 and 53: Nephtys hombergii's spatial distrib
- Page 54 and 55: (vii) G. duebeni (ix) % Organic con
- Page 56 and 57: 8m survey - spatial patterns Figure
- Page 58 and 59: (1) P. elegans (iii) L. conchilega
- Page 60 and 61: a) Ts 1.4 0.6 u 0.2 -0.2 1.4 'E5 0.
- Page 62 and 63: 200m 150m 100m 50m (ix) C. edule 56
- Page 64 and 65: 73 ‘a• el 1.4 (ix) G. duebeni 1
- Page 66 and 67: DISCUSSION The main aims of this st
- Page 68 and 69: formed patches less than 1m2 and th
- Page 70 and 71: stutchbutyi, at Wirroa island, New
- Page 72 and 73: exhibited by the tube-building poly
- Page 74 and 75: CHAPTER 3 THE POPULATION STRUCTURE
- Page 76 and 77: Asexual reproduction by fragmentati
- Page 78 and 79: METHODS Survey design - It has been
- Page 80 and 81: RESULTS The species abundances in e
- Page 82 and 83: corresponds to 44 setigers using Eq
- Page 84 and 85: 1 0000000 00 rg 0 00 d- - Xauanbau
- Page 86 and 87: Reproductive activity of Pygospio e
- Page 88 and 89: P. elegans larvae at Drum Sands hav
- Page 90 and 91: Pygospio elegans showed great seaso
- Page 92 and 93: Previous studies have produced simi
- Page 96 and 97: abundance are highly seasonal, were
- Page 98 and 99: CHAPTER 4 THE EFFECTS OF MACROALGAL
- Page 100 and 101: studies may have been completely di
- Page 102 and 103: METHODS Study site - The exact posi
- Page 104 and 105: 1 C N W 4----111" 1.5m 2 NW C Contr
- Page 106 and 107: sediment sampling, together with re
- Page 108 and 109: RESULTS Species abundances - The me
- Page 110 and 111: ; 15 35 — 30 — 25 — 10 — 5
- Page 112 and 113: statistical difference from net plo
- Page 114 and 115: Pygospio elegans size distribution
- Page 116 and 117: used, approximately equivalent to t
- Page 118 and 119: artefacts associated with the metho
- Page 120 and 121: present in high numbers around sewa
- Page 122 and 123: lack, hydrogen sulphide-smelling se
- Page 124 and 125: CHAPTER 5 THE EFFECTS OF MACROALGAL
- Page 126 and 127: METHODS Survey design - During late
- Page 128 and 129: The sediments could not be sampled
- Page 130 and 131: RESULTS Species abundances - Table
- Page 132 and 133: 90 — 80 — "-e-' 70 — 60 — 4
- Page 134 and 135: 35 — *** 30 25 — 1.) = .-c‘l
- Page 136 and 137: Pygospio elegans size distributions
- Page 138 and 139: which is difficult to compare with
- Page 140 and 141: eason why some invertebrates showed
- Page 142 and 143: This study did not set out to expli
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