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Other theories have been postulated for the observed community differences within tube-beds. Woodin (1978, 1981) suggested that the observed effects result from the 'refuge' provided by the high numbers of tubes. Both predation and local disturbances have been shown to be important structuring forces for soft-bottom infauna (Reise, 1978; Woodin, 1978; Zajac and Whitlatch, 1982a; Ambrose, 1984; Thrush, 1988; Hall et al., 1993) and Woodin (1978) suggested that refuges may be created by organisms whose structures buffer the impact of physiological stress or inhibit the access of predators. Woodin (1976) viewed soft-bottom community structuring from the perspective of 'functional group' interactions in which 'tube- dweller', `bioturbator' and `suspension-feeder' functional groups should each be unfavourable to each other. This 'functional group' interaction hypothesis implied that the maintenance of discrete dense assemblages in infaunal systems was due to interactions between established infaunal individuals and settling larvae. Therefore, a tube-building species would be expected to exhibit increased densities within an assemblage of other tube-builders by this interaction, whilst bioturbators and suspension-feeders would be expected to exhibit lower densities. The main tube-building polychaete species forming dense tube-beds in northern European shallow waters tend to be either Lanice conchilega or Pygospio elegans. Several studies such as Eagle (1973, 1975), Carey (1982) and Ragnarsson (1996) have described the fauna within L. conchilega tube-beds, however, relatively few studies have focused on the beds formed by P. elegans (e.g., Dupont, 1975; Morgan, 1997). The dense tube-beds at Drum Sands allowed an investigation of the effects of this tube-builder on an intertidal sandflat. Specifically, the differences in the biology and the sediments within P. elegans patches and non-patch areas of sand were studied. 203
METHODS The sampling for the investigation described in Chapter 6 was carried out at the same time and within the same patch and non-patch plots as those for the present study. However, a further survey was also carried out in August 1998 for this study. Faunal sampling methods, sample treatment and sorting, faunal identification and statistical analyses for this chapter are as described in Chapter 6. While sediment variables were only measured on one occasion for the investigation described in Chapter 6, the same variables were measured at each sample date for the present study, i.e., April, August and December 1997, and August 1998. In addition to the size measurements of P. elegans, individuals of other species with mean abundances above 20 per core in patch and non-patch samples were measured. P. elegans and C. capitata were measured as 5th setiger and 3rd setiger widths respectively. Small bivalves were measured as shell width using a microscope (x20 magnification) fitted with an eyepiece graticule while larger ones were measured with a pair of calipers. 204
- 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 182 and 183: In April, when P. elegans larval av
- 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 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
- Page 232 and 233: 3NP 6NP 4NP 1 NP 5NP 2NP : 3P 1P 6P
- Page 234 and 235: 4P 3P 5P 5NP 6P 2P 1P Figure 8.8: T
- Page 236 and 237: Figure 8.10 shows the dendrogram pr
- Page 238 and 239: NP1 NP2 NP2 NP2 NP1 NP1 NP2 NP2 NP2
- Page 240 and 241: Sediment water, organic and silt/cl
- Page 242 and 243: 5 350 — 300 250 200 — ISO — 1
- Page 244 and 245: abundances of P. ciliata had more d
- Page 246 and 247: levels of silt/clay and organics. S
- Page 248 and 249: 1973; Noji and Noji, 1991). Competi
- Page 250 and 251: shown to consume up to 68% of a 0-g
- Page 252 and 253: In Chapter 7 the micro-scale spatia
- Page 254 and 255: and positions of patches. Consequen
- Page 256 and 257: epresent those found establishing i
- Page 258 and 259: distribution at the micro-scale. Ad
- Page 260 and 261: provide a rich food source for deme
- Page 262 and 263: Armonies W., 1988. Active emergence
- Page 264 and 265: Cha M.W., in prep. Macroalgal mats
- Page 266 and 267: Dobbs F.C. and Vozarik J.M., 1983.
- Page 268 and 269: Flach E.C., 1996. The influence of
Other theories have been postulated for the observed community differences within<br />
tube-beds. Woodin (1978, 1981) suggested that the observed effects result from the<br />
'refuge' provided by the high numbers of tubes. Both predation and local<br />
disturbances have been shown to be important structuring forces for soft-bottom<br />
infauna (Reise, 1978; Woodin, 1978; Zajac and Whitlatch, 1982a; Ambrose, 1984;<br />
Thrush, 1988; Hall et al., 1993) and Woodin (1978) suggested that refuges may be<br />
created by organisms whose structures buffer the impact of physiological stress or<br />
inhibit the access of predators. Woodin (1976) viewed soft-bottom community<br />
structuring from the perspective of 'functional group' interactions in which 'tube-<br />
dweller', `bioturbator' and `suspension-feeder' functional groups should each be<br />
unfavourable to each other. This 'functional group' interaction hypothesis implied<br />
that the maintenance of discrete dense assemblages in infaunal systems was due to<br />
interactions between established infaunal individuals and settling larvae. Therefore, a<br />
tube-building species would be expected to exhibit increased densities within an<br />
assemblage of other tube-builders by this interaction, whilst bioturbators and<br />
suspension-feeders would be expected to exhibit lower densities.<br />
The main tube-building polychaete species forming dense tube-beds in northern<br />
European shallow waters tend to be either Lanice conchilega or Pygospio elegans.<br />
Several studies such as Eagle (1973, 1975), Carey (1982) and Ragnarsson (1996) have<br />
described the fauna within L. conchilega tube-beds, however, relatively few studies<br />
have focused on the beds formed by P. elegans (e.g., Dupont, 1975; Morgan, 1997).<br />
The dense tube-beds at Drum Sands allowed an investigation of the effects of this<br />
tube-builder on an intertidal sandflat. Specifically, the differences in the biology and<br />
the sediments within P. elegans patches and non-patch areas of sand were studied.<br />
203