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•ins 11.. I on a tidal Hat in the Wadden Sea after cold and mild winters. J. exp. mm. Biol. Ecol. 153: 97-113. Beukema J.J 1992a. Dj namics of juvenile shrimp Crangon crangon in a tidal Hal nursery of the Wadden Sea alter mild and cold winters. Mar Ecol. Prog. Ser 83: Is7-I65. Beukema J.J. 1992b. Expected changes in the Wadden Sea benthos in a warmer world: lessons from periods with mild winters. Neth. J. Sea Res 30:73-79. Beukema JJ. 1993a. Increased mortality in alternative bivalve prey during a period when the tidal Hals of the Dutch Wadden Sea were devoid of Mussels. Nelh. J. Sea Res. 31: 395-406. Beukema JJ. 1993b. Successive changes m distribution patterns as an adaptive strategy in the bivalve Macoma balthica (l-.l in Ihe Wadden Sea. Helgolander Meeresunters 47: 2K7 304 Beukema JJ. & W. de Bruin 1977. Seasonal changes in dry weighl and chemical composition of the soli parts ol ihe lelhnid bivalve Macoma balthica in ihe Dutch Wadden Sea. Neth, J. Sea Res. 11:42-55. Beukema JJ. & G.C. Cadee 1991. Growlh rates of the bivalve Ma coma balthica in the Wadden Sea during a period ol eutrophi- cation: relationships with concentrations of [vlagic diatoms and flagellates. Mar. Ecol Prog. Ser. 68: 249-256. Beukema JJ. & R. Dekker 1995. Dynamics and growlh ol a recent invader into European coastal waters: the American raZCi clam. Ensis directus. J. mar. biol. Ass. U.K. 75: 351-362. Beukema JJ. A: M. Desprez 1986 Single and dual annual grow ing seasons in ihe tellinid bivalve Macoma balthica (L.I J. exp. mar. Biol. Ecol. 102: 35-45. Beukema JJ. & K. Essink 1986 Common patterns in ihe fluctua tions of macro/.oobcnihic species living al different places on tidal Hats ii) the Wadden Sea. Hydrobiologia 142:-199 207 Beukema J.J. & B.W. Median 1985. Latitudinal variation in linear growlh and oilier shell characterisiics oi Macoma balthica Mar. Biol. 90; 27-33. Beukema JJ. & J. de Vlas 1979. Population parameters nl the Lug worm. Arenicola marina, living on tidal Hats in the Dutch Wadden Sea. Neth. J. Sea Res. 13: 331-353. Beukema J.J.. W. de Bruin & JJ.M. Jansen 1978. Biomass and species richness of the macrohenthic animals living on the tidal Hats of the Dutch Wadden Sea long-term changes during a period wilh mild w inters. Nelh. J. Sea Res. 12: 58-77. Beukema J.J.. E. Knol & G.C. Cadee 1985. Effects of temperature on the lengih of the annual growing season in the tellinid bivalve Macoma balthica (L.) living on lidal Hals in the Dutch Wadden Sea. J. exp. mar. Biol. Ecol. 90: 129-144 BeiikemaJJ.. K. Fssink H Miehaehs A- I ZwaTO 1993 Ycar-lo- yeni variability in macrohenlhic biomass on tidal Hats in Ihe Wadden Sea: how predictable is ilus lood source for birds'.' Nelh. J. Sea Res. 31:319-330. Bijlsma R.G. 1990. Predation by large falcons on wintering waders on ihc Banc d'Arguin. Mauritania. Ardea 78: 75-82. REFERENCES 370 Birklund J. 1977. Biomass. growlh and prixluclion of the amphipcxl Coropliiiini iiisidiosuin Craw lord, and preliininarv notes 00 Corophium votutator (Pallas). Ophelia 16: 187-203. Blomert A-M.. M. Engelmoer & D. Logemann 1983 Voedsel- ivecologie van de Scholeksler op het Frie.se wad. Repon Rijks dienst Usselmeerpolders. Lelysiad. Blundon J.A. & V.S. Kennedy 1982a. Mechanical and behavioural aspects of Blue Crab, Callinectes sapidus (Rathbun), predaiion cm Chesapeake Bay bivalves. J. exp. mar. Biol. Ecol. 65:47-65. Blundon J.A. & V.S. Kennedy 1982b. Refuges for infaunal bivalves from Blue Crab. Callinectes sapidus (Ralhbun). predaiion in Chesapeake Bay. J. exp. mar. Biol. Ecol. 65: 67-81. Boales IS. 1988 Foraging and social behaviour of the Oysier- caicher Haematopus ostralegus in relation to diet specializa tion. Thesis University of Exeler. Exeter. Bi ..des J.S. & J.D. Goss-Custard 1989. Foraging behav iour of Oys- lercaichers Haematopus ostralegus during a did switch from worms Nereis diversicolor to clams Scrobicularia plana. Can. I. Tool 67:2225-2231, Boates J.S. & J.D. Goss-Custard 1992. loraging behaviour of Oys- leicatchers Huemalopiis ostralegus specializing on different species of prey. Can. J. Zool. 70: 2398-2404. li. .ales J.S. & P.C. Smith 1979 Length-weight relationships, energy content and the effects of predaiion on Corophium votutator (Pallas) (Cnisiacea: Amphipoda). Proc. Nova SCOII.I Insl Sci 29: 489-499. Boates J.S. & P.C. Smith 1989. Crawling behaviour of the amphipod Coropliiiini votutator and loraging by Semipalmated Sand pipers. Calidris pus-ilia. Can. J. Zool. 67: 45 Boere G.C. & CJ. Sunt 1981. Knot iCululns canutus L). In: CJ, Smil ft WJ. Wolff (eds.) Birds ol the Wadden Sea: 136-145. Balkema. Rotterdam. Bohnecke G. & G. Dietrich 1951 Monalskarten der Oberflachen- lemperatur I'm die Nord- und Ostsee und die angrenzen- den Gewasser. Deutsches Hydrographisches Institut, Ham burg. Bonsdorff E. & R. Wenne 1989. A comparison of condilion indices • i Macoma balihica (L.I from ihe northern and southern Baltic- Sea. Nelh. J. Sea Res. 23: 45-55. Boulding E.G. I9S4. Crab resisianl features ol shells ol burrowing bivalves decreasing vulnerability by increasing handling time J exp. mar. Biol. Ecol. 74: 201-223 Boyden CR. 1971. A comparative study of the reproductive cycles of Ihe cockles Cerastmlerma edule and ('. viaticum. I. mar. biol. Ass. U.K. 51: 605-622 Brafield A.E. 1964. The oxygen content of interstitial waler ir, sand) shores. J Arum Ecol. 33:97-116. Brafield A.E. & G.E. Newell 1961. The behaviour of Macoma balthica (L). I mar. hid. Ass. U.K. 41: 81-87. BreumO. 1970. Stimulation of burrowing activity hy wave action in some manne bivalves. Ophelia 8: 197-207.
BreyT. 1989. Der Einfluss physikaliseher und biologischer Faktoren auf Struktur und Dynamik der subhtoralen M. balthica- Gemeinschafl der Kieler Buchl. Ber. Inst. Meeresk. Kiel 186: 1-248. Brey T. 1991. Interactions in soft bottom benthic communities: quantitative aspects of behaviour in the surface deposit feeders Pygospio elegan.s tPolychaeta) and Macoma balthica (Bivalvia). Helgolander Mecresunlers. 45: 301-316. BreyT. H. Rumohr& S. Ankar 1988 Energy content of macrohenthic invertebraies: general conversion factors from weight to energy. J. exp. mar. Biol. Ecol. I 17: 271-278. Brousseau D.J. 1978. Population dynamics of the Soft-shell Clam Mya armaria. Mar. Biol. 50: 63-71. Brousseau D.J. 1979. Analysis of grow ih rate in Mya arenaria using Ihe Von Bertalanffy equation. Mar. Biol. 51:221 -227. Brown J.S.. B.P. Kotler. RJ. Smith & W.O. Wirtz II 1988. The effects of owl predation on Ihe foraging behav UH ..I lie leromyid rodents. Oecologia (Berl.) 76: 408-415. Brown R.A. & R.J. O'Connor 1974. Some observations on Ihe tela lionships between Oystercatchers Haematopus ostralegus L. and Cockles Cardium eduli I. in Strangford Lough. Irish Naturalist 18:73-80. Bryan G.W. & L.G, Hummerstone I97S. Heavy metals in the burrowing bivalve Scrobicularia pinna from contaminated and uncontaminated estuaries. J. mar. biol. Ass. U.K. 58: 401-419. Bryant DM. & J. Leng 1975. Feeding distribution and behaviour ..I Shelduck in relation lo food supply. Wildfowl 26: 20-30. Bryant D.M. & P. Tatner 1991. Intraspecilic variation in avian energy expenditure: correlates and constraints. Ibis 133: 236- 245. Bunskoeke EJ.. 1988. Overde Ilucluaiies van de prooikeus van de Scholekster (Haematopus ostralegus) in het Uuedsei/oen 1986op Schiermonnikoog. Student report Zoological Laboratory. University of Groningen. Groningen. Bunski>ekc E.J.. BJ. Ens. J.B. Hulscher & S J.dc Vlas 1996. Why do Oy stercatchers Haematopus oslralegus switch from feeding on Bailie Tellin Macoma balihica to feeding on Ihe Ragworm Nereis diversicolor during the breeding season'.' Ardea 84: in press. Burton PJ.K. 1974. Feeding and ihe feeding apparatus in waders. British Museum. London. Busines J.O. & K.E. Erikslad 1990. Size selection of common Mussel-. Msiilus edulis, by Common Eiders. Somateria mollissinni energ] maximization or shell weighl minimization .'Can. J. Zool. 68: 2280-2283 Buxton N.E. & CM. Young 1981. The food of ihe Shelduck in nonh-easi Scotland. Bird Study 28: 41-48. Caddy J.F. 1967. Maturation of gametes and spawning in Macoma balthica iL.). Can. J. Zool. 45:955-965. CadeeG.C. 1976. Sediment reworking by Arenicola marina on tidal Hats in Ihe Dutch Wadden Sea. Neth. J. Sea Res. 10: 440-460 REFERENCES *r-*i Cain DJ. & S.N. Luoma 1986. Effect of seasonally changing tissue weight on trace metal concentrations in ihe bivalve Macoma balihica in San Francisco Bay. Mar. Ecol. Prog. Ser. 28: 209- 217. Calder W.A. Ill 1974. Consequence of body size for avian energetics. In: R.A. Paymer (ed.i Avian energetics: 86-151 Nlltfall Ornithological Club. Cambridge. Mass. Camphuysen C.J., BJ. Ens, D. Heg. J.B. Hulscher. J. van der Meer & CJ. Smit 19%. Oyslercalchei• Haematopus ostralegus winter mortality in Ihe Netherlands: the effect of severe weather and food supply Ardea 84: in press. Cayford J.T. |9SSa. A field test ol die accuracy of estimating prey size seleciion in Oystercatdiers from recovered mussel shells. Wader Study Group Bull. 54: 29-32 Cayford J.T. 1988b. The foraging behaviour of Oystercatcher (Haematopus oslralegus) feeding on MusselstMstilu.s edulis). Thesis University of Exeter, Exeter. Cayford J.T. & J.D. Goss-Cuslard 1990. Seasonal changes in the size seleciion ol Mussels. Msiilus edulis, by Oystercatchers, Huemalopiis oslralegus: an optimaliiy approach. Alum. Behav 40: 609-624. Cerri R.D. & D.F. Fraser 1983. Predaiion and risk in foraging minnows: balancing conflicting demands. Am. Nat. 121: 552-561. Chambers M.R. & H. Milne 1975a. Theproduclion ot Macoma balthica (I... I in ihe Ythan estuary. Estuar. csti. mar. Sci. 3: 443-455. Chambers M.R. & H. Milne 1975b. Life cycle and production ol Nereis diversicolor OF Miiller in the Ythan Estuary. Scotland. I -iii.n csil. mar. Sci. 3: 133-144. Chambers M.R. & H. Milne 1979. Seasonal variation in the condilion of some inienidal invertebrates of the Ythan estuary. Scolland Estuar, estl. mar. Sci. 8: 411-419. Chapman G. & G.E. Newell 1956. The role of Ihe body fluid in the movement ol soft-bodied invertebrates II. The extension of ihe siphons of Mya arenaria and Scrobicularia plana (da Costa). Proc R SOC B. 145: 564-580. Charnov E.L. 1976. Optimal foraging: attack strategy of a manlid. Am. Nat. 110: 141-151. Cole H.A. 1956. A preliminary study of growth-rate in Cockles (Cardium edule L.) in relation to commercial exploitation. J. Cons. perm, int Explor. Mer. 22: 77-90. Colijn F. 1982. Lighl absorption in [he waters of the Ems-Dollard Estuary and its consequences for the growth of phytoplankton and microphytobenlhos. Neth. J. Sea Res. 15: 196-216. Colijn F. & V.N. de Jonge 1984. Primary production of microphytobenlhos in The Ems-Dollard Estuary. Mar. Ecol. Progr. Ser 14 185-196. Collins M.D.. S.A. Ward & A.F.G. Dixon 1981. Handling lime and ihe functional response of Aphelinus ilumisoni. a predator and parasite of ihe aphid Drepanosiphum pluianoidis. J. Anim. Ecol. 50: 479-487. Commito J.A. 1982. Effects of Lunatia hews predation on Ihe pop-
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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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Page 324 and 325: VERSATILITY OF CURLEWS FEEDING ON N
Page 326 and 327: total time spent al the surface. Th
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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
Page 341 and 342: SAMENVATTING 345
Page 343 and 344: Voorgeschiedenis In 1960 verscheen
Page 345 and 346: maken. Als gebied A een grotere voe
Page 347 and 348: 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
Page 353 and 354: het toen niet gemakkelijk hebben ge
Page 355 and 356: doende nonnetjes van 10 tot 15 mm l
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: Allen RL. 1983. Feeding behaviour o
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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