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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
- 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
- Page 144 and 145: This reliance upon the early establ
- Page 146 and 147: CHAPTER 6 INITIAL COLONISATION OF D
- Page 148 and 149: esulting community at any stage of
- Page 150 and 151: ambient sediment had been removed.
- Page 152 and 153: emoved since they were the only tax
- Page 154 and 155: All statistics were performed using
- Page 156 and 157: RESULTS Univariate analysis of spec
- Page 158 and 159: 3.5 3 5 2 11 5 1 0.5 0 40 35 Ca 30
- Page 160 and 161: of non-patch areas (Figure 6.3(vi))
- Page 162 and 163: the individuals colonising patch az
- Page 164 and 165: Multivariate analysis of community
- Page 166 and 167: Month Sample statistic (Global R) N
- Page 168 and 169: 2NP 3NP 4NP .•,, 6NP 5NP 6P 1NP i
- Page 170 and 171: Figure 6.8: Two-dimensional MDS ord
- Page 172 and 173: - - 5P ... 4P . 6P • .‘2NP 1NP
- Page 174 and 175: I 50. 1 60. 70. 80. 90. 100. BRAY-C
- Page 176 and 177: 'P2-AZ P3-AZ N2-AZ .- - - " .„ ..
- Page 178 and 179: o • o -o + 350 — 300 = 250 7 g
- Page 180 and 181: The importance of the ambient commu
- Page 184 and 185: not only for errant polychaetes, bu
- Page 186 and 187: observed in this study. How crucial
- Page 188 and 189: Micro-scale spatial patterns of mac
- Page 190 and 191: METHODS Experimental design - A pre
- Page 192 and 193: study. These individuals would not
- Page 194 and 195: RESULTS Pilot survey - The pilot su
- Page 196 and 197: Transect survey - Micro-scale patte
- Page 198 and 199: Month v:m ratio pattern Id pattern
- Page 200 and 201: (i) March 1997, replicate 1 -iAlmiA
- Page 202 and 203: (xix) October 1997, replicate 1 (ra
- Page 204 and 205: The new recruits were only sufficie
- Page 206 and 207: The results of correlation analyses
- Page 208 and 209: cf.) . crt N ,—, Cr) C,1 ,—, Cr
- Page 210 and 211: 1.2 -0.4 "a 0.8 > (i) % Water conte
- Page 212 and 213: examine the micro-scale spatial pat
- Page 214 and 215: Invertebrate larvae, those of polyc
- Page 216 and 217: laboratory observations are needed
- Page 218 and 219: CHAPTER 8 THE FAUNAL COMMUNITIES OF
- Page 220 and 221: Other theories have been postulated
- Page 222 and 223: RESULTS Univariate analysis of spec
- Page 224 and 225: -T. g 80 g 50 40 30 20 10 (i) Adult
- Page 226 and 227: in significant differences in size
- Page 228 and 229: 8.2). This was mainly because of th
- Page 230 and 231: 120 100 80 60 - 40 20 0. cn1 c.n (i
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