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50 [® 4.0 •3 3.0 n a •». V TT 0 D DC a T3 c D D 0 B £ 2.0 0 **L4r**-«. B **L4r**-«. in » T ^---a 9 °*8« * O) E "—" f 10 7 i' 0.032x"° 0.001 a* 0.8 Im 047 ra I 0.6 a T • On ° - • «° ° » • "T • • PREV PROFITABILITY AND INTAKE RATE 0.4 1 1 1 1 20 30 40 50 60 shell length (mm) ;® •?* a a 01a 0 T T O X*' - s ^ a a a 0 n^ar aa ° rsSgr* • •*, . •_s**r*W*"' . ' ,^-*tn° rfa'ajPa y=0.092n«*» . « ^ **. ' p
prey species, using parallel slopes for all species (exponent = 0.375) added 31% to the 18';; of the explained variance that could be attributed to prey weighl using the single regression line (exponent = Q 204) in Fig. 11B. On the assumption that all the regression lines for the armoured prey species were not parallel. 5.0 4.0 ® in 2.0 Ol E CO 3.0 - * S 1.0 ID *= 0.8 0 6 0 ••• • y.0.157x0"5 p f\l • • • • 1 ' 1 ' 7 8 9 10 20 30 shell length (mm) 40 the explained variance increased again, but from only 49% to 56%. We therefore conclude that (11 in cadi species, the intake rate increases with prey weight. (21 ihe rate of increase differed between soft-bodied and armoured prey species, (3) the rate of increase did not differ much among the four soft-bodied and among the ;® • • y=0.423x°3» P'OOOI 1=0.65 • • • • \ / • • 3v ' • 1 . 1 1 11 l_ l . , ,1 5 10 50 100 500 1000 prey weight (mg) Kij*. 13. Hie intake rale as a function of A. shell length and B. average weight of Cerastoderma taken. Sources: Brown & O'Cotmoi (1974), Davidson (1967). Drinnan (1957). Hns aal (1996b & 1996c). Exo. Smit & Zvvans (unpubl. i. Goss-Custard 11977), Hulscher 11976 & unpubl.). Leopold ei al. (1989). Meire (1996b). Sutherland (1982b & 1982c). 20 30 40 shell length (mm) 20 40 100 400 prey weight (mg) Fig. 14. The intake late as a funclion of A. shell length and B. average weighl of Scrobicularia laken. Sources: Blomert et al. (1983). Boates & Goss-Custard I I9X9|. Habckolte' (1987). Hulscher et al. (unpubl.). Wanink & Zwarts (1985 & 1996). Zwarts (unpubl.). 193 •• 40 60 80 100 prey weight (mg) Pig, 15, The intake rale as a lunciion ol ihe average weight of Nereis taken. Sources: Boates & Goss-Custard (1989). Bunskoeke et al. (1996). Durell el al. (1996 & unpubl). Hulscher (unpubl. >. Triplet (1989). •
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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
Page 137 and 138: 100 200 300 prey density (n-m-2) Fi
Page 139 and 140: Table I. Results of five one way an
Page 141 and 142: Table 2. Results of eight iwo-wav a
Page 143 and 144: OPTIMAL FORAGING AND THE FUNCTIONAL
Page 145 and 146: prey density II III Fig. 12. Freque
Page 147 and 148: OPTIMAL FORAGING AND THE FUNCTIONAL
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 181 and 182: 20 30 40 50 shell length (mm) PREY
Page 185 and 186: Nereis Arenicola tipuia earthwo'm M
Page 187: take rate. We might therefore expec
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
Page 199 and 200: situation arrived in the western pa
Page 203 and 204: • ' • 14 PREY PROFITABILITY AND
Page 205 and 206: Notes lo appendix: PREY PROFITABILI
Page 207: Chapter 10 WHY OYSTERCATCHERS HAEMA
Page 210 and 211: 2 4 6 8 10 time on feeding area (h)
Page 212 and 213: 80- 60- o 80 60- 40- 20- INTAKE RAT
Page 214 and 215: est of bod\ behind: an estimated 22
Page 216 and 217: INTAKE RATE AND PROCESSING RATE IN
Page 218 and 219: Wanink 1993). The energy content of
Page 220 and 221: (7) Age All studies dealt with adul
Page 222 and 223: irds over long periods. As an examp
Page 224 and 225: Discussion There is no difference i
Page 226 and 227: comparison between the weight of ih
Page 228 and 229: . 1', L ! •
Page 230 and 231: Introduction PREDICTING SEASONAL AN
Page 232 and 233: PREDICTING SEASONAL AND ANNUAL FLUC
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PREDICTING SEASONAL AND ANNUAL FLUC
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1000 r 1 II" 1 - r^Jan'80 PREDICTIN
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IOO BO 60 40 20 0 PREDICTING SEASON
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WHY KNOT TAKE MEDIUM-SIZED MACOMA W
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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
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50 100- s s I — f = S. plana, n=2
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area is twice as large as the touch
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10 15 lower size threshold (mm) Fig
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lime (Hughes 1979). This would furt
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WHY KNOT TAKE MEDIUM-SIZED MACOMA T
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Chapter 13 ANNUAL AND SEASONAL VARI
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VARIATION IN FOOD SUPPLY OF KNOT AN
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Two sites (N and M in Fig. 2) were
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20 30 VARIATION IN FOOD SUPPLY OF K
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20 l i r'l ' i •o j^y^T n ^ ' " ^
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July ' Aug ' Sept Fig. 9. Proportio
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available. There must, however, be
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Chapter 14 SEASONAL TREND IN BURROW
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BURROWING AND FEEDING IN NEREIS SEA
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Table 2. Results of a 2-way analysi
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June 1981 June 1982 is * N.MUD •
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8 10 I 12 JZ 5. -g 1*» •e o i 16
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6 - n=!080 5 • g 4 CP > i 3 CO CD
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1985) and Curlew (/wails ,v, Lsseli
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Chapter 15 VERSATILITY OF MALE CURL
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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
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VERSATILITY OF CURLEWS FEEDING ON N
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total time spent al the surface. Th
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2 4 . • 2.2 • 30 .-.2.0 to Q. 1
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PREY DEPLETION BY OYSTERCATCHER AND
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the rule: the observed lower limit
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to locate the clams after they had
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Curlew, by bury ing some hundreds o
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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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