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abundances of P. ciliata had more diverse meiofaunal and macrofaunal communities compared with areas of low abundances, while Reise (1983b) found that the presence of dense assemblages of P. elegans promoted the abundance of small benthic organisms by approximately 40%. Morgan (1997), using a correlation approach, indicated that the majority of the most common taxa in the Baie de Somme, France, e.g., Eteone longa, Hediste diversicolor and Cerastoderma edule, were significantly positively correlated with P. elegans abundances. Many studies endeavouring to elucidate the mechanism by which high numbers of tube-builders affect infaunal community structure have focused on the way in which recolonisation is effected (e.g., Gallagher et al., 1983; Trueblood, 1991). Resident adults may influence colonists in many ways determined by their feeding mode and modifications to the sediments and hydrodynamics (Thrush et al., 1992). However, experimental studies on such mechanisms have proved equivocal. Since recruitment is usually assayed some time after settlement and metamorphosis (Bachelet, 1990) the actual patterns of larval settlement are often obscured (Hadfield, 1986). Consequently, studies have failed to assess the mechanisms by which tube-builders affect recolonisation since it is inherently difficult to distinguish between differential settlement and differential mortality of larvae (Woodin, 1986). Larvae of some marine benthic invertebrate species have been experimentally shown to actively select settlement sites using certain cues (Scheltema, 1974; Woodin, 1986; Butman et al., 1988a; Pawlik and Butman, 1993; Hsieh, 1994). However, it is likely that in the field, water flow is greater than the swimming speeds of larvae and instead, larvae are transported as passive particles and deposited via passive entrainment (Hannan, 1984; Butman, 1987; Butman et al., 1988b). Eckman (1983) predicted that velocity of near-bed flow through plastic straws (seagrass mimics) would be reduced to such an extent that suspended particles, such as larvae and meiofauna, would be passively deposited. His results supported this prediction although his experimental design was flawed by replication of only one of his treatments, his defaunated control. Later experiments have incorporated replication of the defaunated controls but the results have been inconclusive due to a number of factors including a plot size too small to allow the formation of a fully developed boundary layer (Kern and Taghon, 227
1986) and low level of replication resulting in too low a statistical power (Ragnarrson, 1996). The present study was observational and not intended to determine the mechanisms responsible for any differences in abundances between patch and non-patch communities. However, the increased silt/clay fraction and higher meiofaunal abundances in patches may have been due to increased passive deposition due to the reduction in velocity of near-bed flow. In the same way, passive larval entrainment may be responsible for some of the observed differences in this study. This was observed even though tube-building spionids have been shown to ingest settling bivalve larvae (Breese and Phibbs, 1972; Daro and Polk, 1973). Once an individual of a species had colonised a P. elegans patch, the physico-chemical effects of the tubes possibly concurred to provide an increased food supply in the form of the flourishing microbial and meiofaunal communities observed in this study. Furthermore, the resistance to shearing forces provided by the beds may have allowed a dense community by virtue of individuals not being 'swept' away (Morgan, 1997). Increased abundances of meiofauna (43%) were observed by Reise (1983b) in P. elegans patches compared to areas lacking the spionid, whilst a similar increase has been documented for beds of another spionid, Polydora ciliata, by Noji (1994), who suggested that the meiofauna were utilising the worms faecal pellets as a food source. The presence of C. volutator in patches, while almost completely absent outside patches, probably resulted from active habitat selection of adults to areas of increased silt/clay fraction and more stabilised sediments. This amphipod attains particularly high densities in muddy sediments with high numbers of diatoms (Lawrie, 1996). Its almost exclusive existence within patches suggests some sort of functional group interaction (sensu Woodin, 1976) for this tube-building, deposit-feeding amphipod. Similarly, Reise (1978) noticed C. volutator beds harbouring large numbers of P. elegans in the Wadden Sea. Sediment differences between P. elegans patches and non-patch areas. Sediment analyses indicated that although sediment water content was the same for patch and non-patch areas, the former consistently contained significantly increased 228
- Page 194 and 195: RESULTS Pilot survey - The pilot su
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- Page 200 and 201: (i) March 1997, replicate 1 -iAlmiA
- Page 202 and 203: (xix) October 1997, replicate 1 (ra
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- Page 206 and 207: The results of correlation analyses
- Page 208 and 209: cf.) . crt N ,—, Cr) C,1 ,—, Cr
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- 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
- 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 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
- Page 270 and 271: Hall SI, Raffaeni D.G., Basford DJ.
- Page 272 and 273: Keckler D., 1997. Surfer for Window
- Page 274 and 275: Levin L.A. and Creed E.L., 1986. Ef
- Page 276 and 277: Mileikovsky S.A., 1971. Types of la
- Page 278 and 279: Ong B. and Krishnan S., 1995. Chang
- Page 280 and 281: Raffaelli D.G., Hildrew A.G. and Gi
- Page 282 and 283: Scheltema R.S., 1974. Biological in
- Page 284 and 285: Soulsby P.G., Lowthion D., Houston
- Page 286 and 287: McArdle B.H., Morrisey D., Schneide
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- Page 290 and 291: Zajac R.N. and Whitlatch R.B., 1982
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abundances of P. ciliata had more diverse meiofaunal and macrofaunal communities<br />
compared with areas of low abundances, while Reise (1983b) found that the presence<br />
of dense assemblages of P. elegans promoted the abundance of small benthic<br />
organisms by approximately 40%. Morgan (1997), using a correlation approach,<br />
indicated that the majority of the most common taxa in the Baie de Somme, France,<br />
e.g., Eteone longa, Hediste diversicolor and Cerastoderma edule, were significantly<br />
positively correlated with P. elegans abundances.<br />
Many studies endeavouring to elucidate the mechanism by which high numbers of<br />
tube-builders affect infaunal community structure have focused on the way in which<br />
recolonisation is effected (e.g., Gallagher et al., 1983; Trueblood, 1991). Resident<br />
adults may influence colonists in many ways determined by their feeding mode and<br />
modifications to the sediments and hydrodynamics (Thrush et al., 1992). However,<br />
experimental studies on such mechanisms have proved equivocal. Since recruitment<br />
is usually assayed some time after settlement and metamorphosis (Bachelet, 1990) the<br />
actual patterns of larval settlement are often obscured (Hadfield, 1986).<br />
Consequently, studies have failed to assess the mechanisms by which tube-builders<br />
affect recolonisation since it is inherently difficult to distinguish between differential<br />
settlement and differential mortality of larvae (Woodin, 1986).<br />
Larvae of some marine benthic invertebrate species have been experimentally shown<br />
to actively select settlement sites using certain cues (Scheltema, 1974; Woodin, 1986;<br />
Butman et al., 1988a; Pawlik and Butman, 1993; Hsieh, 1994). However, it is likely<br />
that in the field, water flow is greater than the swimming speeds of larvae and instead,<br />
larvae are transported as passive particles and deposited via passive entrainment<br />
(Hannan, 1984; Butman, 1987; Butman et al., 1988b). Eckman (1983) predicted that<br />
velocity of near-bed flow through plastic straws (seagrass mimics) would be reduced<br />
to such an extent that suspended particles, such as larvae and meiofauna, would be<br />
passively deposited. His results supported this prediction although his experimental<br />
design was flawed by replication of only one of his treatments, his defaunated control.<br />
Later experiments have incorporated replication of the defaunated controls but the<br />
results have been inconclusive due to a number of factors including a plot size too<br />
small to allow the formation of a fully developed boundary layer (Kern and Taghon,<br />
227