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\* w * ,!i 4- > .«? «? •r; V". VARIATION IN FOOD SUPPLY OF KNOT L$L-, ' /; fr *=« w V 6' M >»-fil* .A V' v 4V* / ' V •' •it 1 • ( ' 1 Si V "i J ?• '%'^ v - * s) . , I he Mud Snail may occur in huge densities at the substrate and hence seems to be an easy prey for Knol. Knot prefer, however, mediuni-si/ed. iliin-slielled bivalves. 294 1 t
20 l i r'l ' i •o j^y^T n ^ ' " ^ 11. 20 - 11) \\ r\ 0 n VARIATION IN FOOD SUPPLY OF KNOT Macoma balihica Cerastodeniia edule 1111111 1111111111111111111111111111111111111 JJMMJSN'JMMjSN'JMMJSN'jMMJSN 1 Fig. 5. Growth of five bivalve species at site SJ after spatfall in sumuicr 1979 (1983 for St nihil ularia plana). The si/e classes vulnerable lo prcdaiion h\ Knol (See fable 2) are indicaied (horizontal grey fieldl. as well as ihe period during which Knot were present (vertical grey bars). Average sizes were calculated from the length lie queue) distribution* in the monthly samples (see 'Methods' i ten years. The contribution of the 1979 year class of Macoma lo the biomass harvestable by Knot is evident: the harvestable food supply was highest alter four growing seasons and decreased thereafter because an increasing proportion had passed the critical size threshold of 16 mm. On some occasions. Macoma was the only prey species harvestable hv Knot for example in late summer of 1981 and 1983. Though Cerastoderma were available in 1983. Knot ignored them, perhaps due to the thick shell (Zwarts & Blomen 1992). Macoma musl therefore have been the staple food for Knot that passed through ihe study area in late summer, unless the Mud Snail Peringia ulvae was taken as an alterna- 295 tive. This species usually occurred ai densities of 40 000 to 50 000 nv'. though the summer of 1979 was an exception, when only low densities of 1000 m : occuned (Zwarts 19881*0 EJowevet direel observations and faecal analysis showed that Peringia were nol taken in 1979. 1981. 1983 or 1985. even though the) were abundant in three of ihe four years. In fact, the majority of Peringia in the study area were too small « 2 mm) to be profitable food items for Knot (Zwarts & Blomert 1992). The distribution paiiern of Knol over ihe feeding area was another indication lhat Knot did not feed on Peringia. This prey was superabundant at the sites situated above mean sea level and in areas where the clay content of the substrate was above 10% (Zwarts 1988b). However. Knot ignored these areas and were usually found on sites below mean sea level where the clay content of the substrate was less than This resulted in a negative relationship between density of Knot and that of Peringia. Considering all this information together, the conclusion is that Macoma was the main, and sometimes the only, prey for Knot in the study area. For thai reason, local and sea sonal variations only in Macoma are described below. 1977 78 79 80 81 82 83 84 85 86 tig. 6. Biomass (g AFDW m -i ol live bivalve species M sik- \ m August from 1977 to 19X6. A selection has been made for the range of si/e classes barvulihlc b) Knol Calidris , annua (see fable 2). No correction has been made for depth distribution of the prey.
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WADERS AND THEIR ESTUARINE FOOD SUP
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plia^ohi. WADERS AND THEIR ESTUARI
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Waders and their estuarine food sup
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WADERS AND THEIR ESTUARINE FOOD SUP
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figuren: Dick Visser omslagfoto: Ja
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15 Versatility of male curlews (Num
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That science is making progress, ma
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INTRODUCTION The remnants of brushw
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When hcnlhic bivalves extend llien
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the burrow depths and on the fracti
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INTRODUCTION Ms,i invest 409 of (he
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able lo sw itch 10 oiher prey which
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Long-term, broadly-based research c
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Chapter 1 SEASONAL VARIATION IN BOD
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SEASONAL VARIATION IN BODY WEIGHT O
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Tahle 1 Weight loss ('•- ± SE) m
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i 60 50 40 30 20 10 0 • INFESTED
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240- SEASONAL VARIATION IN BODY WEI
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20 10 0 -10 -20h 2 3 4 5 6 7 8 9 se
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a E 400 - S. plana 35 mm 320 240 16
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Chapter 2 HOW THE FOOD SUPPLY HARVE
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FOOD SUPPLY HARVESTABLE BY WADERS H
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Methods The study sites were situat
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less than that of bivalves, partly
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I Fig. 3). Peak condition was reach
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total biomass since most of those t
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on the basis of the preceding and t
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However, for obvious reasons, we ma
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prey. A decrease in the prey densit
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Tahle 2. The intake rate ol < lyste
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* ' % -. . ; 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
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Chapter 3 BURYING DEPTH OF THE BENT
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DEPTH AND SIPHON CROPPING IN SCROBI
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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
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Chapter 4 SIPHON SIZE AND BURYING D
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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
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4 Q) Si •a a. 16- 20- A predator:
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prey risk for an animal al a depth
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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
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surface in die containers was varie
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50 OOF 310.00 I 5.00 ^ 1.00 3=" 0.5
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Tahle 1. Macoma and Scrobicularia.
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siphon would not have to reach the
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known siphon weight and burying dep
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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
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Table 2. Results of eight iwo-wav a
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prey density II III Fig. 12. Freque
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PREY SIZE SELECTION AND INTAKE RATE
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Predicted 'passive size selection'
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lig. 2. Larger Mussels are less ava
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Fig. 5. Scrobicularia plana. Size c
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and maiiv are too thick-shelled to
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The measurements of handling time i
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1 2 3 4 5 6 7 probing depth (cm PRE
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valves may vary by a factor of two
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optimal foraging model, the minimum
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their gut is full? Do they reduce t
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PREY PROFITABILITY AND INTAKE RATE
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catchers to take only certain prey
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licult error of estimate arose if p
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Counts of feeding and non-feeding b
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20 30 40 50 shell length (mm) PREY
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Nereis Arenicola tipuia earthwo'm M
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take rate. We might therefore expec
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prey species, using parallel slopes
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5.0 4.0 1-3.0 a & • % * • * •
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time during the feeding period. As
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winter. Similarly, handling time in
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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
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• ' • 14 PREY PROFITABILITY AND
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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
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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
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PREDICTING SEASONAL AND ANNUAL FLUC
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Page 240 and 241: 1000 r 1 II" 1 - r^Jan'80 PREDICTIN
Page 242 and 243: PREDICTING SEASONAL AND ANNUAL FLUC
Page 252 and 253: IOO BO 60 40 20 0 PREDICTING SEASON
Page 265 and 266: WHY KNOT TAKE MEDIUM-SIZED MACOMA W
Page 267 and 268: in Zwarts & Esselink 1989). The len
Page 269 and 270: shell length (mm) FIR. 3. Peringia.
Page 271 and 272: 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: 20 30 VARIATION IN FOOD SUPPLY OF K
Page 293 and 294: July ' Aug ' Sept Fig. 9. Proportio
Page 295 and 296: 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,
Page 316 and 317: 0.6 0.8 1.0 1.2 1.4 1.6 jaw lengih
Page 318 and 319: Na oi both Nrieck Fig. 7. Numenius
Page 320 and 321: 6 8 10 12 14 16 18 worm length (cm)
Page 322 and 323: too (Fig. 11 A). Indeed, the averag
Page 324 and 325: VERSATILITY OF CURLEWS FEEDING ON N
Page 326 and 327: 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
Page 335 and 336: 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
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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
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kunnen opzuigen. of diep leven en z
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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
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was dan ook dank/ij de inzet van al
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REFERENCES 367
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Allen RL. 1983. Feeding behaviour o
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BreyT. 1989. Der Einfluss physikali
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latie tol chironoiiiiilen en de wai
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huch der Vogel Milteleuropas, Band
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llolmann II. & H. Huerschelmann 196
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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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