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t CONTENTS DECLARATION. . ii ABSTRACT . . iii ACKNOWLEDGEMENTS. . v CONTENTS . . vi LIST OF FIGURES . x LIST OF TABLES .Xiii CHAPTER 1. INTRODUCTION. Background . . 1 The present study . 6 Pygospio elegans . 8 The study site . . 10 CHAPTER 2. THE SMALL- TO MESO-SCALE SPATIAL DISTRIBUTION OF . . 15 MARINE BENTHIC INVERTEBRATES ON DRUM SANDS, WITH PARTICULAR REFERENCE TO PYGOSPIO ELEGANS Introduction . . 15 Methods . . 19 Scales of spatial variability - Transect survey. . 19 Survey design. . 19 Data analyses . . 21 Pattern analysis - Grid surveys . 22 Survey design . . 22 Data analyses . . 24 Results. . . 27 Scales of spatial variability - Transect survey. . 27 Pattern analysis - Grid surveys . 31 lm survey - spatial patterns . 36 8m survey - spatial patterns . 42 40m surveys - spatial patterns. . 43 Discussion . 51 CHAPTER 3. THE POPULATION STRUCTURE AND REPRODUCTIVE . . 59 BIOLOGY OF PYGOSPIO ELEGANS ON DRUM SANDS. Introduction . . 59 Methods . . 63 Survey design . . 63 Data analyses . . 64 vi
Results. . 65 Size distribution of Pygospio elegans. . 65 Sex ratio of Pygospio elegans. . . 70 Reproductive activity of Pygospio elegans . 71 Discussion . . 74 CHAPTER 4. THE EFFECTS OF MACROALGAL COVER ON THE SPATIAL . 83 DISTRIBUTION OF MACROBENTHIC INVERTEBRATES: AN EXPERIMENTAL STUDY. Introduction . . 83 Methods . . 87 Study site . . 87 Experimental design . 87 Data analyses . . 91 Results. . 93 Species abundances . 93 Sediment water, organic and silt/clay contents and . 94 granulometry Redox potentials . . 97 Pygospio elegans size distribution . 99 Algal biomass. . 100 Discussion . . 102 CHAPTER 5. THE EFFECTS OF MACROALGAL COVER ON THE SPATIAL . 109 DISTRIBUTION OF MACROBENTHIC INVERTEBRATES: A SURVEY APPROACH Introduction . . 109 Methods . . 111 Survey design . . 111 Data analyses . . 114 Results. . 115 Species abundances . . 115 Sediment water, organic and silt/clay contents and . 118 granulometry Redox potentials . . . 118 Pygospio elegans size distribution . 121 Algal biomass. . 122 Discussion . . 124 vii
Page 1 and 2: AN INVESTIGATION INTO THE PROCESSES
Page 3 and 4: ABSTRACT The spionid polychaete Pyg
Page 5: ACKNOWLEDGEMENTS I am indebted to m
Page 9 and 10: CHAPTER 9. GENERAL DISCUSSION . . 2
Page 11 and 12: Figure 5.4 Figure 5.5 Figure 6.1 Fi
Page 13 and 14: LIST OF TABLES Table 2.1 Statistica
Page 15 and 16: BACKGROUND CHAPTER 1 INTRODUCTION A
Page 17 and 18: The scales of observation, or the s
Page 19 and 20: systematic sampling design to inves
Page 21 and 22: aised sediment within an otherwise
Page 23 and 24: Fauchald and Jumars (1979) describe
Page 25: Dalmeny House and sewage discharged
Page 28 and 29: E 7— E 6-ac. MHWS MHWN t co 4 —
Page 30 and 31: variance (TTLQV) techniques (see Lu
Page 32 and 33: analysis using Moran's and Geary's
Page 34 and 35: Holme and McIntyre (1984). Percenta
Page 36 and 37: Pattern Analysis - Grid Surveys Sur
Page 38 and 39: 57 64 1=1 0 0 0 0 0 0 0 O 0 0 0 0 0
Page 40 and 41: Maps produced by kriging and other
Page 42 and 43: 200 180 1160 140 100 1-3 80 g 60 c.
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
Page 48 and 49: " v : m pattern Id pattern Ip patte
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 94 and 95:
The sole reliance on a planktonic m
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
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 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
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(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
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
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
Page 288 and 289:
Wharfe J.R., 1977. An ecological su
Page 290 and 291:
Zajac R.N. and Whitlatch R.B., 1982
Page 292 and 293:
- C'e) oc0000mooF,o•-ooNtn-coo-00
Page 294 and 295:
APPENDIX 1.2: SPECIES ABUNDANCES FR
Page 296 and 297:
ON In co In ° ken n00000N---0g0N-.
Page 298 and 299:
o m000eno,-0-. 0 0000en ,-. 0.-00 o
Page 300 and 301:
APPENDIX 2: SPECIES ABUNDANCES FROM
Page 302 and 303:
0.- CV N - , .- CI E.104 ..r cv g.,
Page 304 and 305:
cv en 0 a .- ,- .- .- g c', ,- ,- g
Page 306 and 307:
PI — — — Pi n — — - R., .
Page 308 and 309:
P . en .2 Co in ,.. ne .- cq . -- -
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