waders and their estuarine food supplies - Vlaams Instituut voor de ...

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$Wetenschap - \Nijverheid- Handel$

figuren: Dick Visser omslagfoto: Jan van de Kant lulu's \un landschappen, bodemdieren en vogels: Jan van tic Kam overige loin's: Jan H. Wanink en Leo Zwarts vogeltekeningen: Jos Zwarts DTP en drukwerh Evers Litho & Druk codrdinatie productie: Henk Bos
CONTENTS Introduction 9 1 Seasonal variation in body weight of the bivalves Macoma halihica. Scrobicularia plana, Mya arenaria and Cerastodema edule in the Dutch Wadden Sea 25 2 How the food supply harvestable by waders in the Wadden Sea depends on the variation in energy density, body weight, biomass. burying depth and behaviour of tidal-flat invertebrates. 45 3 Burying depth of the benthic bivalve Scrobicularia plana (da Costa) in relation to siphon-cropping. 83 4 Siphon si/e and burying depth in deposit- and suspension-feeding benthic bivalves. 95 5 Feeding radius, burying depth and siphon length of Macoma halthica and Scnibicularia plana. 111 6 The maerobenlhos fraction accessible to waders often represents marginal prey. 127 7 Doev an optimally foraging oystercatcher obey the functional response' 137 8 Prey size selection and intake rate. 153 9 Causes nl variation in prey profitability and its consequences lor the intake rate of 0\ stercatehers llticnuiiopiis oslralegus. 173 10 Why Oystercatcher Haeinatopiis nstralegus cannot meet their daily energy requirements in a single low water period. 211 11 Predicting seasonal and annual fluctuations in the local exploition of different prey by ()\ steicatcher Haematopus oslralegus: a ten year study in the Wadden Sea. 231 12 Why knot Calklris eanutus take medium-sized Macoma bnliltica when six prey species are available. 267 13 Annual and seasonal variation in the food supply harvestable by knot Calidrii caiuittis staging in the Wadden Sea in late summer. 287 14 Seasonal trend in burrow depth and tidal variation in surface feeding of Xereis diverslcolor. 301
Page 1 and 2: WADERS AND THEIR ESTUARINE FOOD SUP
Page 3 and 4: plia^ohi. WADERS AND THEIR ESTUARI
Page 5 and 6: Waders and their estuarine food sup
Page 7 and 8: WADERS AND THEIR ESTUARINE FOOD SUP
Page 12 and 13: 15 Versatility of male curlews (Num
Page 15 and 16: That science is making progress, ma
Page 17 and 18: INTRODUCTION The remnants of brushw
Page 19 and 20: When hcnlhic bivalves extend llien
Page 21 and 22: the burrow depths and on the fracti
Page 23 and 24: INTRODUCTION Ms,i invest 409 of (he
Page 25 and 26: able lo sw itch 10 oiher prey which
Page 27 and 28: Long-term, broadly-based research c
Page 29 and 30: Chapter 1 SEASONAL VARIATION IN BOD
Page 31 and 32: SEASONAL VARIATION IN BODY WEIGHT O
Page 33 and 34: Tahle 1 Weight loss ('•- ± SE) m
Page 39 and 40: i 60 50 40 30 20 10 0 • INFESTED
Page 41 and 42: 240- SEASONAL VARIATION IN BODY WEI
Page 43 and 44: 20 10 0 -10 -20h 2 3 4 5 6 7 8 9 se
Page 45 and 46: a E 400 - S. plana 35 mm 320 240 16
Page 49 and 50: Chapter 2 HOW THE FOOD SUPPLY HARVE
Page 51 and 52: FOOD SUPPLY HARVESTABLE BY WADERS H
Page 53 and 54: Methods The study sites were situat
Page 55 and 56: less than that of bivalves, partly
Page 57 and 58: I Fig. 3). Peak condition was reach
Page 59 and 60:
total biomass since most of those t
Page 61 and 62:
on the basis of the preceding and t
Page 63 and 64:
However, for obvious reasons, we ma
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prey. A decrease in the prey densit
Page 67 and 68:
Tahle 2. The intake rate ol < lyste
Page 69 and 70:
* ' % -. . ; a X - . - * • ^ •
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(Zwarts & Wanink 1989). In quiet we
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general law relating handling time
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nearly twice as much time (0.79 s).
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4 5 6 7 8 9 size class of Corophium
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and annually. Second, the lower siz
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Prey switching Waders feeding on ti
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autumn and spring, and the contrary
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where they switch to surface-living
Page 87 and 88:
Chapter 3 BURYING DEPTH OF THE BENT
Page 89 and 90:
DEPTH AND SIPHON CROPPING IN SCROBI
Page 91 and 92:
I DEPTH AND SIPHON CROPPING IN SCRO
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DEPTH AND SIPHON CROPPING IN SCROBI
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Table 3. Three-Way analysis Of vari
Page 97:
Chapter 4 SIPHON SIZE AND BURYING D
Page 100 and 101:
15 20 size (mm) Fig. 3. Cerasioderm
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10 size (mm) SIPHON SIZE AND DEPTH
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o, 7 01 5 | * CL 5- SIPHON SIZE AND
Page 106 and 107:
4 Q) Si •a a. 16- 20- A predator:
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prey risk for an animal al a depth
Page 110 and 111:
Table 6. Size al which there is a m
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* ' 1 /-/ "».-^*«C
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face and so expose themselves to a
Page 116 and 117:
surface in die containers was varie
Page 118 and 119:
50 OOF 310.00 I 5.00 ^ 1.00 3=" 0.5
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Tahle 1. Macoma and Scrobicularia.
Page 122 and 123:
siphon would not have to reach the
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known siphon weight and burying dep
Page 127 and 128:
ACCESSIBLE PREY ARE OFTEN IN POOR C
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1 2 3 4 burying depth (cm) Kin. I.
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I—1 • • : : : : ! - ! -|-r 0
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Chapter 7 DOES AN OPTIMALLY FORAGIN
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OPTIMAL FORAGING AND THE FUNCTIONAL
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100 200 300 prey density (n-m-2) Fi
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Table I. Results of five one way an
Page 141 and 142:
Table 2. Results of eight iwo-wav a
Page 143 and 144:
Page 145 and 146:
prey density II III Fig. 12. Freque
Page 147 and 148:
Page 149:
PREY SIZE SELECTION AND INTAKE RATE
Page 152 and 153:
Predicted 'passive size selection'
Page 154 and 155:
lig. 2. Larger Mussels are less ava
Page 156 and 157:
Fig. 5. Scrobicularia plana. Size c
Page 158 and 159:
and maiiv are too thick-shelled to
Page 160 and 161:
The measurements of handling time i
Page 162 and 163:
1 2 3 4 5 6 7 probing depth (cm PRE
Page 164 and 165:
valves may vary by a factor of two
Page 166 and 167:
optimal foraging model, the minimum
Page 168 and 169:
their gut is full? Do they reduce t
Page 171 and 172:
PREY PROFITABILITY AND INTAKE RATE
Page 173 and 174:
catchers to take only certain prey
Page 175 and 176:
licult error of estimate arose if p
Page 177 and 178:
Counts of feeding and non-feeding b
Page 179 and 180:
Page 181 and 182:
20 30 40 50 shell length (mm) PREY
Page 183 and 184:
Page 185 and 186:
Nereis Arenicola tipuia earthwo'm M
Page 187 and 188:
take rate. We might therefore expec
Page 189 and 190:
prey species, using parallel slopes
Page 191 and 192:
5.0 4.0 1-3.0 a & • % * • * •
Page 193 and 194:
time during the feeding period. As
Page 195 and 196:
winter. Similarly, handling time in
Page 197 and 198:
5.0 - f-4.0 S 3.0 in I 20 a 2 a | 1
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situation arrived in the western pa
Page 201 and 202:
Page 203 and 204:
• ' • 14 PREY PROFITABILITY AND
Page 205 and 206:
Notes lo appendix: PREY PROFITABILI
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Chapter 10 WHY OYSTERCATCHERS HAEMA
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2 4 6 8 10 time on feeding area (h)
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80- 60- o 80 60- 40- 20- INTAKE RAT
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est of bod\ behind: an estimated 22
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INTAKE RATE AND PROCESSING RATE IN
Page 218 and 219:
Wanink 1993). The energy content of
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(7) Age All studies dealt with adul
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irds over long periods. As an examp
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Discussion There is no difference i
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comparison between the weight of ih
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. 1', L ! •
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Introduction PREDICTING SEASONAL AN
Page 232 and 233:
PREDICTING SEASONAL AND ANNUAL FLUC
Page 234 and 235:
Page 236 and 237:
Page 238 and 239:
Page 240 and 241:
1000 r 1 II" 1 - r^Jan'80 PREDICTIN
Page 242 and 243:
Page 244 and 245:
Page 246 and 247:
Page 248 and 249:
Page 250 and 251:
Page 252 and 253:
IOO BO 60 40 20 0 PREDICTING SEASON
Page 254 and 255:
Page 256 and 257:
Page 258 and 259:
Page 260 and 261:
Page 262 and 263:
Page 265 and 266:
WHY KNOT TAKE MEDIUM-SIZED MACOMA W
Page 267 and 268:
in Zwarts & Esselink 1989). The len
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shell length (mm) FIR. 3. Peringia.
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fused, specimens larger than this w
Page 273 and 274:
50 100- s s I — f = S. plana, n=2
Page 275 and 276:
area is twice as large as the touch
Page 277 and 278:
10 15 lower size threshold (mm) Fig
Page 279 and 280:
lime (Hughes 1979). This would furt
Page 281 and 282:
WHY KNOT TAKE MEDIUM-SIZED MACOMA T
Page 283 and 284:
Chapter 13 ANNUAL AND SEASONAL VARI
Page 285 and 286:
VARIATION IN FOOD SUPPLY OF KNOT AN
Page 287 and 288:
Two sites (N and M in Fig. 2) were
Page 289 and 290:
20 30 VARIATION IN FOOD SUPPLY OF K
Page 291 and 292:
20 l i r'l ' i •o j^y^T n ^ ' " ^
Page 293 and 294:
July ' Aug ' Sept Fig. 9. Proportio
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available. There must, however, be
Page 297 and 298:
Chapter 14 SEASONAL TREND IN BURROW
Page 299 and 300:
BURROWING AND FEEDING IN NEREIS SEA
Page 301 and 302:
Table 2. Results of a 2-way analysi
Page 303 and 304:
June 1981 June 1982 is * N.MUD •
Page 305 and 306:
8 10 I 12 JZ 5. -g 1*» •e o i 16
Page 307 and 308:
6 - n=!080 5 • g 4 CP > i 3 CO CD
Page 309 and 310:
1985) and Curlew (/wails ,v, Lsseli
Page 311:
Chapter 15 VERSATILITY OF MALE CURL
Page 314 and 315:
Smid! 1951. Muus 1967, Wolff 1973,
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0.6 0.8 1.0 1.2 1.4 1.6 jaw lengih
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Na oi both Nrieck Fig. 7. Numenius
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6 8 10 12 14 16 18 worm length (cm)
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too (Fig. 11 A). Indeed, the averag
Page 324 and 325:
VERSATILITY OF CURLEWS FEEDING ON N
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total time spent al the surface. Th
Page 328 and 329:
2 4 . • 2.2 • 30 .-.2.0 to Q. 1
Page 331 and 332:
PREY DEPLETION BY OYSTERCATCHER AND
Page 333 and 334:
the rule: the observed lower limit
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to locate the clams after they had
Page 337 and 338:
Curlew, by bury ing some hundreds o
Page 339 and 340:
PREY DEPLETION BY OYSTERCATCHER AND
Page 341 and 342:
SAMENVATTING 345
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Voorgeschiedenis In 1960 verscheen
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maken. Als gebied A een grotere voe
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omdat ze nog vrijwel niets wegen. l
Page 349 and 350:
kunnen opzuigen. of diep leven en z
Page 351 and 352:
van de uitgerekte sifo over het wad
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het toen niet gemakkelijk hebben ge
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doende nonnetjes van 10 tot 15 mm l
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aangevoerde voedsel en de wulp past
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zijn dan twee jaar en wulpen geen s
Page 361 and 362:
was dan ook dank/ij de inzet van al
Page 363 and 364:
REFERENCES 367
Page 365 and 366:
Allen RL. 1983. Feeding behaviour o
Page 367 and 368:
BreyT. 1989. Der Einfluss physikali
Page 369 and 370:
latie tol chironoiiiiilen en de wai
Page 371 and 372:
huch der Vogel Milteleuropas, Band
Page 373 and 374:
llolmann II. & H. Huerschelmann 196
Page 375 and 376:
l-cndrem D.W. 1984. Flocking, feedi
Page 377 and 378:
Pekkarinen M. 1984. Regeneration of
Page 379 and 380:
lion of Mussels Mytilus edulis hy O
Page 381 and 382:
lulal Dal esiuanes: a comparison of
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