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ulaiion dynamics of Mya arenaria and Macoma balthica in Maine. USA. Mar. Biol. 69: 187-193. Cook R.M. & BJ. Cockrell 1978. Predator ingestion rate and Us bearing on feeding time and the theory ol optimal diels J Anim. Ecol. 47: 529-547. Craeymeersch J A.. P.M.J. Herman & P.M. Meire 1986. Secondary production of an intertidal mussel (Mytilus edulis 1..I popula tion in ihe Eastern Scheldt (S.W. Netherlands). Hydrobiologia 133: 107-115. Ciam A. & S.M. Evans 198(1 Stability and lability in the evolution of behaviour in Nereid polychaeles. Anim. Behav. 28: 483- 490. Cramp S. & K.E.L. Simmons (eds.) 1983. The birds of the western I'.ilacarctic. Vol. III. Oxford University Press. Oxford. Cranford P.J.. D.L. Peer& D.C. Gordon 1985 Population dynamics and production of Macoma balthica in Cumberland Basin and Shcpody Bay, Bay of Fundy. Nelh. J. Sea Res. 19: 135- 146 Crisp D.J. 1984. Energy flow measurements. In: N.A Holme ."c A.D. Mclntyre (eds.) Methods for the study of marine benthos. Blackwell Scientific Publications. Oxford. Curio E. 1976. The ethology of predation. Spnnger Vcrlag, Berlin. Curtis D.J.. C.G. Galbraith. J.C. Smyth & D.B.A. Thompson 1985 Seasonal variations in prey selection by estuarine Black- headed Gulls (Larus ridibundus). Estuar. Csil. Shelf Sci. 21: 75-90. Daan S. & P. Koene 1981. On the timing ol loraging flights by Oys tercatchers, Haematopus ostralegus, on lidal Hats. Neth. J. Sea Res. 15: 1-22. Daan S.. D. Masman Sr. A. Groenewold 1990. Avian basal metabolic rales: their association with body composition and energy ex penditure in nature. Am. J. Physiol. 259: 333-340. I l.iles R.R 1950. The reproduction and larval development af Nereis diversicolor Of. Miiller. J. mar. biol. Ass. U.K. 24: 321-360. Dales R.R 1951. An annual history of a population of Nereis diver sicolor OV. Miiller Biol. Bull. 101: 131-137. Dales R.P. & G. Y. Kennedy 1954. On the diverse colours of Nereis diversicolor. J. mar. biol. Ass. U.K. 33:699-708. Dankers N. & J J. Beukema 1984. Distributional patterns of macro- zoobenthic species in relation lo some environmental factors In: W.J. Wol II led i Ecology of the Wadden Sea, Vol. I. part 4: 69-103. Balkema. Rotterdam. Dankers N. & M.A. Binsbergen 1984. Zeeduizendpoten (Nereis di versicolor) trekken ook. levende Natuur 85: 14-15. Dare PJ. 1975. Settlement, growth and production of Ihe Mussel. Msiilus edulis I... in Moreeambe Bay. England. Fish. Invest Lond (Ser. Ill 28: 1-25. Dare P.J. 1977. Seasonal changes in body-weight of oystercatchers Haematopus ostralegus. Ibis 119: 494-506. Dare PJ. & D.B. Edwards 1975. Seasonal changes in flesh weighl and biochemical composition of Mussels (Mytilus edulis Inn REFERENCES 5 77 the Conwy estuary, North Wales. J exp mar. Biol. Ecol. 18: 89-97. Dare PJ. & AJ. Mercer 1973. Foods of ihe oystercatcher in More eambe Bay. Lancashire. Bird Study 20: 173-184 Dauvin J. & M Joncoun 1989. Fneigy values of marine benlhic in vertebrates from ihe western English Channel. J. mar. biol. \.- I K 69 589-595. Davey J.T. & CL. George 1986. I actors in ihc distribution of intertidal, estuarine polychaeles: a field experiment wilh Nereis (Hediste) diversicolor and Nephtys hombergii in the Tamar al Plymouth. Estuar. Cstl. Shelf Sci. 22: 603-618, Davidson D 1978. Experimental tests ol the optimal diet in two social insects. Behav. Ecol.Sociobiol. 4: 35-41. Davidson P.E. 1967. A sludy of the Oystercatcher {.Haematopus oslralegus L.i in relation to the fishery for Cockles (Cardium eduk L.) in the Burry Inlet. South Wales Fish. Invest 1-ond (Ser. 11)25: 1-28. Davidson P.E. 1971. Some foods taken by waders in Moreeambe Bay. Lancashire. Bird Study 18: 177-186. Davies N.B. 1977. Prey selection and social behaviour in wagtails (Aves Mii.icillidae). J. Anim. Ecol. 46: 37-57. Dekinga A. & T. Piersma 1993. Reconstructing diet composition on Ihe basis of faeces in a mollusc-eating wader, the Knol Calidris canutus. Bird Sludy 40: 144-156. Dekker R. 1979. Numbers, growlh. biomass and production of or ganic and calcareous matter of Hydrobia ulvae (Gastropoda: 1'rosobranchiai in the western Dutch Wadden Sea. NIOZ Report 1979-15 Netherlands Institute for Sea Research, Texel. Desprez M., J-P. Ducrotoy & B. Sylvand 1986. Fluctuations na- turelles et evolution artificielle des biocenoses macrozooben- lluques mteilid.iles de trois estuaires des coles Irani,.uses de la Manehe. Hydrobiologia 142: 249-270. Desprez M.. G. Bachelet. J J. Beukema. J-P. Ducrotoy. K Essink. J Marchand. H. Michaelis. B. Robineau & J.G. Wilson 1991. Dynainique des populations de Macoma balihica (Ll dans les estuaires des Nord-Ouesi de I'Europe: premiere synthesc. In: M. Elliott & J-P. Ducrotoy (eds.i Estuaries and coasts spatial and temporal intercomparisons: 159-165. Olsen & Olsen. Fre- denshore Diamond J.M.. W.H. Karasov. D. Phan & F.L. Carpenter 1986. Digestive physiology is a determinant of foraging boul fre quency in hummingbirds. Nature 68: 225-252. Dick W.J.A.. M.W Pienkowski. M. Waltner & C.D.T Minton 1976. Distribution and geographical origins of Knot Calidris canutus in Europe and Africa. Ardea 64: 22-47. Dill L.M. & A.H.G. Fraser 1984. Risk nl predaiion and feeding behavior of juvenile Coho Salmon (Oncorhsnchus ki.siitch) Behav. Ecol. Sociobiol. 16: 65-71. Dirksen S.. M. van Wouwe & H. Smit 1992. Watervogels in het zourwatergetijdengebied Ventjagersplaten (Haringvliet) in re-
latie tol chironoiiiiilen en de waierstand. Report RIZA. Dordrecht. Drinnan R.E. 1957. The winter feeding of Ihe Oystcrcatclier (Haematopus ostralegus) on the Edible Cockle (Cardiion edule). J. Anim. Ecol 26: 441-469. Drinnan R.E. 1958a. The wintci feeding of the tSjrstsreatcher (Haematopus ostralegus) on the Edible Mussel (Mytilus edulis) III ihe Conway estuary, North Wales i-ish Invest Load. 22: 1-15. Drinnan RE. 1958b. Observations on the feeding of the Oyster- catchei in captivity. Brit. Birds 51: 139-149. Ducrotoy J-P.. H. Rybarczyk. J. Souprayen. G. Bachelet. J.J. Beukema. M. Desprez. J. Dorics. K Essink, J. Guillou. H Miehaelis. B. Sylvand, J.G. Wilson. B. Elkai'm & F. Ibanez 1991. A comparison of the population dynamics of the Cockle (Cerastoderma edule. L.) in North-Westem Europe. In: M. Elliott & J-P Ducrotoy (eds.) Estuaries and coasts: spatial and temporal intercomparisons: 173-184. Olsen & Olsen. Fredensborg, Dugan PJ 1981 The importance of nocturnal feeding in shorebirds: a consequence of increased invertebrate prev activ nv In: N.V. Jones & W.J. Wolff (eds.) Feeding and survival Strategies ol estuarine organisms: 251-260. Plenum Press. New York. Durell S.E.A. le V. dil & J.D. Goss-Custard 1984. Prey selection within a size-class ol Mussels. Msiilus edulis. hy Oyster catchers. Haeinaiopus ostralegus, Anim Behav. 32: 1197- 1203 Durell S.E.A. le V. dil. J.D. Goss-Custard & A. Perez lluii.ido 1996. The efficiency of juvenile Oystercalchers Huemalopiis ostralegus feeding on Ragworms Nereis diversicolor. Ardea SI in press. Earll R. 1975 Temporal variation in the heart activity ol Scrobicularia plana ida Cosun in constant and tidal conditions .1 exp mar. Biol. Ecol 19: 2s7-274. ds D.C ft J.D Huebrter 1977. Feeding and growth rates of Polinices diiplicaius preying on Af.vo armaria at Barnstable Harbor, Massachusetts EcolOg* ; s 1218-1236. mis R. & J.H. Steele 1968. The ecology ol Ihe 0-groop Plaice and Common Dabs at Loch Ewe I Population and food. J. exp mar. Biol. Ecol. 2: 215-238. Ehlert W 1964. Zur Okologie und Biologic der Eniatirung einiger l.imikolcn-Arten. J. Orn. 105: 1-53.. Eisma D. 1963. Shell-characteristici of Cardium edule L. as indicator of salinity. Neth. J. Sea Res. 2: 493-540. Elliott J.M. 1983. The responses ol Ihe aquatic parasitoidVi.'".''!/"'^ armatus (Hymenoptera Agjiotypidae) to the spatial distribution and density .it n- caddis host Silo pallipes (Trichoptera: Gocridac). J. Anim. Ecol. 52: 315-330. Emerson C.W.. TE. Minchinlon & J. Grant 1988. Population structure, biomass, and inspiration of Mya arenaria L. on temperate sandflat. J. exp. mar. Biol. Ecol. 115: 99-111. REFERENCES 373 Emlen J.M. 1966. The role of time and energy in food preference. Am. Nat. 109:427-435. Engelmoer M. & A-M. Blomert 1985. Broedbiologie van de Kluui lanes de hriese vvadilenkusi seizoen 1983 RIJP report 1985 39Abw. Rijksdiensl voorde Usselmeerpolder-. I dystad EngclmocrM.. T Piersma. W. Altenburg ft R M. Banc d'Arguin. In: P.R. Evans. J.D. Goss-Cuslard & W.G, Hale (eds. i Coasial waders and wildfowl in winter: 293-310. Camhndge Universiiy Press,Cambridge. Ens HI 1982 size -election In imisseMeeding Oystercatchets Wader Study Group Bull. 34: 16-20. Ens BJ. & D. Ailing 1996a. Prey selection of a captive Oystfl catcher Haematopus ostralegus hammering on Mussels Mytilus edulis. Ardea 84: in press. EnsBJ.&D Ailing 1996b The effect of an experimentally created niusselhecl on bird densities and food inrale of ihe Oysiereaicher Haematopus ostralegus. Ardea 84: in press. Ens BJ. & J.T. Cayford 19%. Feeding wilh other (rystocatchers. In: J.D. Goss-Custard (ed.) The Oystercatcher: from individuals to populations: 77-104. Oxford University Press. Oxford. Ens BJ. & J.D. Goss-Cuslard 1984, Interference among Oystercatchers. Haematopus ostralegus, feeding on Mussels. Msiilus edulis. on the Exe estuary. J. Anm 7 17-231. Ens BJ. & L. Zwarts 1980a. Wulpen op het wad van Moddergat. Watervogels 5: 108-120. Ens B.J. ft L. /.wans 1980b leniionaal gedrag bij Wulpen buiten het broedgebied Watervogels 5: 155-169. Ens B.J.. P. Esselink & L. Zwarts 1990. Kleploparasitism as a problem of prey choice: a sludy on mudflat-fceding Cm lews. Nstmenius arquata. Anim. Behav. 39: 219-230. Ens BJ.. M. Kersten. A. Brenninkmeijer & J. B. Hulscher 1992. Termor, quality, parental effort and reproductive success. •: (ercatcher (Haematopus osirulegus). J. Anim. Ecol. 61: 703- 715. Ens B.J.. M. Klaassen ft I. /wans 1901 Flocking and feeding in the fiddler crab 11 . i/ tangeri): prey availability as risk-taking behaviour. Nelh. J. Sea Res. 31: 477 494. Ens. B.J.. EJ Bunskoeke, R Hoekxtm, J.B. Hulscher& SJ. de Vlas 1996a, Prey choice and search speed: why simple opiimality tails IO explain the pre) choice rj*f Oystercatchers Haematopus ostralegus (ceding on Macoma balihica and Nereis diversicolor. Ardea 84: in press Ens B.J.. S. Dirksen. CJ Sunt & EJ. Bunskoeke 1996b. Seasonal change, m size selection and intake rate of Oystercalchers Haematopus oslralegus feeding on Ihe bivalves Msiilus edulis and Cerastoderma edule. Ardea 84: in press Ens B.J.. T. Merck. CJ. Smit & EJ Bunskoeke 1996c. Functional and numerical response of Oystercatchers Haeinaiopus ostralegus on shellfish populations. Ardea 84: in press Erichscn J.T, J.R. Krebs & A.I. Houston 1980. Optimal foraging and crypnc prev J Anim. Ecol. 49: 271-276.
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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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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
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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 and 366: Allen RL. 1983. Feeding behaviour o
Page 367: BreyT. 1989. Der Einfluss physikali
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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