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I'laiicliiliys flesus population in the western Dutch Wadden Sea Mar. Ecol. Prog. Ser. 73: 141-148. Velilliiu/en 111), van & D.W. Phillips 1978. Prey capture by Pisas- i, i brevispinus | \steroidea: Bcbhtodennata) on soil substrate Mar. Biol 48: 89-97. Verkuil Y. A. Koolhaas & J. van der Winden 1993. Wind effects on prey availability: how northward migrating waders use brack ish and hypersaline lagoons in the Sivash. Ukraine. Nelh. J. Sea Res 31:359-374. Verwey J. 1958. Orientation in migrating marine animals and a com parison with thai ofothei migrants. .Arch, neerl Z.KII. 13: 418- 445. \ unsiein R.W. 1979. Predaiion on estuarine infauna: response pat terns of component species Estuaries 2 Visser M & L.J. Reinders 1981. Waiting lime as a new component in functional response models. Neth J. Zool. 31: 315-328. Vlas J. de 1979a. 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Application of random search and REFERENCES 384 optimal diet models to predict when Oystercalchers start ex ploiting a Cohort of the bivalve Sci-ohii ularia plana, isiihiintied I Warwick R.M. & R. Price 1975. Macrolauiia produciion in an esiu- arine mud-flat. J. mar. biol. Ass U.K. 55: 1-18. Weissburg M. 1986 Risky business: on ihc ecological relevance of nsk sensitive Foraging, Oikos46: 261-262. Welham C.V.J. 1994. Flight speeds Ol migrating buds: atesl ol max imum range speed: predictions from three aerodynamic equa tions. Behav. Ecol 5: 1-8. Werner B. 1956. fiber die Winlerwanderung von Arenicola manna I 1I'olychacla sedentanai. Helgolander Meeresunlers. 5: 353- 378. Wemer E.E.. J.E Gilliam. D.J. Hall & G.G. Mitlelhach 198.3. An ex- iviiinenial lest of the effects of predation nsk on habitat use in fish Ecology 64: 1540-1548. Whitfield D.F.. A.D. Evans & P.A. Whitfield 1988. The impact of raptor predaiion on wintering waders. I'roc. 19th intern. Or- nilh. Congr: 674-687. Whitlow G.C. 1986. Energy metabolism. In: PD. Sturdie (ed). Avian Physiology Springer Vei lag. New York, 253-268. Wiersma P. & T. Piersma 1994. Effects ol inierohabiial. Hocking, climate and migratory goal on energy expenditure in the annual cycle of Red Knots Condor 96: 257-279. Wikander P.B. 19811 Biometry and behaviour in Abra nilida (MQller) and A. longicullii.s iScacchii (Bivalvia. Tellinaceal Saisia 65: 255-268. Wilde P.A. W.J. 1975. Influence Of temperature 00 behaviour, energy metabolism, and growth of Macoma balthica (L.) In: H. Barnes led.) Proc. 9th Europ. mar. biol. Symp.: 239 256 Aberdeen University Press. Aberdeen. Wilde P.A.W.J. de & E.M. Berghuis 1978. Laboratory experiments on the spawning of Macoma balihica: Us implication for pro duciion research. In: D.S. McLusky A: A.J Berry leds.) Physi ology and behaviour ol marine organisms: 375-384. Perga mon. Oxford. Wilde P. AAV J. de & E.M. Berghuis 1979a. Cyclic temperature lluc- luaiuins in a tidal mud-flat. In: E. 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Reproductive cycles ol Cerastoderma glaucum (Brugierel and C edule (L.l wilh special reference to the effects ol the 1981-82 severe winler. J. Moll. Stud. 52: 6-14. Yin M.C. & J.H.S. Blaxter 1987. Escape speeds of marine fish lar vae during early development and starvation Mar. Biol. 96: 459-468. Unsen F. 1991. Karaktcrgelallen van de winters vanaf 1707. Zcnit 18:69-73. Yonge CM. 1953. Aspecis ol hie in muddy shores. In. Essays on manne biology: 29-49. Oliver & Boyd. Edinburgh. Zajac R.N. 1985. The effect oi sublethal predaiion on reproduction in the spionid polychaele Polsdora ligni Webster. J. exp. mar. Biol Ecol. 88: 1-19. Zegers P.M. & N.D. Kwinl 1992. Vogeltellingen in hel Nederlandse deel van het Waddengebied 1979-1990. SOVON-rcport 1992/14, SOVON. Bcek-Ubbergen. Zwarts L. 1974. Vogels van het brakke gctijgehicd. Bondsuitgevenj. Amsterdam. /wans L. 1981. Habitat selection and competition in wading buds In: CJ. Smil & W.J. Wolll (eds.) Birds of Ihc Wadden Sea: 271-279. Balkema. Rotterdam. Zwarts L. 1983. Flukluatics in het vleesgewicht van bodemdieren: Fnes-Groningse Wad 1976-1982. Report Rijksdienst voorde Usselmeerpolders. Leiv siad Zwarts L. 1985. The winter exploitation of fiddler crabs Uca tangerl hy waders m Guinea-Bissau. Ardea 73: 3-12. Zwarts L. 1986. Burying deplh of Ihe benthic bivalve Scrobicularia plana Ida Costa) in relation to siphon cropping. J. exp. mar. Biol. Ecol. 101:25-39. Zwarts L. 1988a. Numbers and distribution of coastal waders in Guinea-Bissau. Ardea 76: 42-55. /warts L. 1988b. De bodemfauna van dc Fries-Groningse wad- denkust. Flevobericht 294. Ki'ksdienst voorde Usselmeerpol- v siad Zwarts L. 1990. Increased prey availability drives preinigration by- REFERENCES 385 perphagia in Whimbrels and allows ihem lo leave Ihe Banc d'Arguin Mauritania in time. Ardea 78: 279-300. Zwarts I. 1991. Seasonal variation in body condition of the bivalves Macoma balihica. Scrobicularia tilanu. Msu armaria and Cerastoderma edule in the Dutch Wadden Sea. Neth. J. Sea Res.28 : : i 243 Zwarts L. & A-M. Blomen 1990. Selectivity of Whimbrels feeding on tiddler crabs explained by component specific dige-libih lies. Ardea 78: 193-208. /.warts L & A-M. Blomen 1992. Why Knol Calidris caiiutus lake medium-sized Macoma balthica when si\ prey species are available. Mar. Ecol Prog Ser. 81: 113-128. /wans L. & A-M. Blomert 1996. Daily metabolized energ) intake ol ( ly-Ici catchers Hacinulopiis oslralegus leeding on larvae ol the crane fly Tipula paludosa. Ardea 84: in press Zwarts L. & S. Dirksen 1990. Digestive bottleneck limits the in crease in food intake of Whunbrels preparing for spring migra tion from the Banc d'Arguin. Mauritania. Ardea 78: 257-278. /wans L. & R.H. Dreni 1981 Prey depletion and ihe regulation of predator density: Oystercatchers (Haematopus ostralegus) leeding 00 Mussels (Mytilus edulis). In: N.V. Jones & W.J. Wolff (eds, i Feeding and survival strategics ol 'estuarine organ isms: 193-216. Plenum Press, New York. Zwarts L. & P. Esselink 1989. Versatility of male Curlews (Nume nius arquata) preying upon Nereis dhri rsu olor. deploying con trasting capture modes dependent on prey availability. Mar. Ecol. Prog. Ser. 56: 255-269. Zwarts L. & J. Wanink 1984. How Oystercatchers and Curlew cessively deplete clams. In: PR Evans. J.D Goss-Custard & W.G. Hale (eds i (nasi., id wildfowl in winter 69- 83. Cambridge University Press. Cambridge. Zwarts L. & J. Wanink 1989 Siphon stage and burying depth in de posit- and suspension-feeding benthic bivalves. Mar. Biol. 100: 227-240. Zwarts L. & J.H. Wanink 1991. The inacrohenlhos fraction at hie to waders may represent marginal prey. Oecologia (Berl.) 87:581-587. Zwarts L & J.H. Wanink 1993. 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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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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
Page 331 and 332: PREY DEPLETION BY OYSTERCATCHER AND
Page 333 and 334: the rule: the observed lower limit
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Page 337 and 338: Curlew, by bury ing some hundreds o
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Page 341 and 342: SAMENVATTING 345
Page 343 and 344: Voorgeschiedenis In 1960 verscheen
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Page 349 and 350: kunnen opzuigen. of diep leven en z
Page 351 and 352: van de uitgerekte sifo over het wad
Page 353 and 354: het toen niet gemakkelijk hebben ge
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Page 357 and 358: aangevoerde voedsel en de wulp past
Page 359 and 360: 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: lion of Mussels Mytilus edulis hy O
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