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coasts: spatial and temporal intercomparisons: 307-116. olsen iV i llscn. I redensborg. Kamermans P. & H.J. Huitema 1994 Shrimp (Cnutgon i rangon L.) browsing upon siphon lips inhibits feeding and growlh in the bi valve Macoma balihica (L.). J. exp. mar. Biol. Ecol. 175: 59-75. Kcl-o WE. 1979. Predaiion on Soli-shell Clams. Mya armaria, by the Common Mummichog, I' iinditlits heteroctitus. Estuaries 7 249-254. Kendeigh S.C., Dol'nik V.R. & V.M. Gavrilov 1977. Avian ener getics. In: J. I'liiouski & S.C. Kendeigh ledsi Granivorous birds in ecosystems. Cambridge University Press. Cambridge, 127-204. Kenvvard R.E. 1978. Hawks and doves: factors affecting success and selection in Goshawk attacks on Woodpigeoiis. J Anim Ecol 47: 449-160. Kenvvard R.E & R.M. Sibly 1977. A Woodpigeon {Columlni piilumhiis) feeding preference explained by a digestive bottle neck. J appl. Ecol. 14:815-826, Kenvvard R.E. & R.M. Sibly 1978. Woodpigeon feeding behav iou at brassica sites. A field and laboratory investigation of Wood pigeon feeding behaviour during adaptation and maintenance of a brassica diet Anim. Behav. 26: 778-790. Kersten. M. 1996. Time and energ) budgets ot Oystercatchers Haematopus ostralegus occupying territories ol different qual ity \nlea S4. in piess. Kersten M. & A. Brenninkmcijer 1995. Growth. Hedging success and post-fledging survival of juvenile Oystercatchers Haematopus ostralegus. Ibi- 137: 396-404. Kersten M. & T Piersma 1984. Voedselkeuze cn voedselopname v.m /ilverplevieren Pliisuihs sipiaiamla ill de Waddenzee tij dens de v.H.r en n.i|aarstrek. Limosa 57: 105-111. Kersten M. & T Piersma 1987. High levels ol energy expenditure in shorehirds: metabolic adaptations to an energetically expen sive way of life. Ardea 75: 175-187. Kersten M. & W. Visser 1996a. The rale of food processing in Oys tercatchers: lood intake and energy expenditure constrained by a digestive bottleneck, Funct. Ecol 10: 440-448. Kersten M. & W. Visser 1996b. Food intake of Oystercatcher by day and by night measured with an electronic nesi balance. Ardea 84: in press. Kim Y.S. 1969. Selective feeding on several bivalve molluscs by the starfish, listerias amurensis. Bull. Fac. Fish. Hokkaido Univ. 19:244-249. Kingston P.P. 1974. Sludies on ihe reproductive cycles of Cardium edule and C. glaiicum. Mar. Biol. 28: 317-323. Kirkegaard J.B. 1978. Settling, growth and life span of some poly chaeles from Danish waters Medd. Danm. Fisk. -og Havun- ders.. N.S. 7:447-496. Klaassen M.. M. Kersten & BJ. Ens 1990. Energetic requirements for maintenance and prcmigratory body mass gain of waders wintering in Africa. Ardea 78: 209-220 REFERENCES 378 Kleel H.I... P. Tydeman & K. Essink 1982. Overde invloed van hel al dan met losprepareren van het vlees op de nauwkeurig- heid van hel biomassagclal bi| Ivveekleppige schelpdieren. Report Rijksinstiluul voor Zuivcring van Afvalwater, Groinn .nen Klein Breteler W.C.M. 1975. Rood consumption, growth and eneigy niclnholisni of juvenile Shore Crabs, Carcinus maenns. Neth. J. Sea Res. 9: 255-272. Koene P. 1978. De Scholekster: uaiilalscttcctcn op de voedselop name. Student report Zoological l-aboratory. University of Groningen. Groningen Kramer D.L.. D. Manley & R. Bourgeois 1983 The effect ol'respi ratory mode and oxygen concentration on the risk ol aerial pre daiion in fishes Can. J. Zool. 61: 653-665. Kiel's J.R. 1978. Optimal foraging: decision rules for predators. In JR. Krebs & N B Ilavies (eds.i Behavioural ecology an evo lutionary approach: 23-63. Blackvvell. Oxford. Krebs J.R. 1980. Optima] foraging, predaiion risk and temtory de- lenee. Ardea 68: 83-90. Krebs J.R. & A. Kacelmk 1991 Decision-making. In: J.R Krebs iv N.B Davie's (eds.) Behavioural ecology an evolutionary ap proach: 105-136. Blackvvell. Oxford. Krebs JR.. D.W. Stephens & W.J. Sulherland 1983. Perspectives in optimal loraging. In: GA. Clark (ed.i Perspectives in orniihol ogy: 165 221. Cambridge University Press. Cambridge Kristensen E. 1981. Direct measurement of ventilation and oxygen uptake in three species ot mbtculous polychaeles (Nereis spp. I J. Comp. Physiol 145:45-50. Kristensen I. 1957. Differences in density and growth in a cockle population in the Dutch Wadden Sea. Arch. Necrl. Zool. 12: 351-453. Kilhl H. 1951. Studien tlberdie Klallmuscliel Mya arenaria 3. Das Junggui. Arch Fischereiwiss. 6: 33-44. Kiihl H. 1981. The Sandgaper. Mya arenaria. In: N. Dankers. II Ktihl & W.J. Wolff (eds i Invertebrates ot the Wadden Sea: 118-119. Balkema. Roilerdani. Kuipers B.R. 1977. On ihe ecology of juvenile Plaice on a tidal flat of Ihe Wadden Sea Neth J. Sea Res. 11: 56-91. I.ael IJR de 1985. Dominance and ami-predator behaviour of Greal Tits Parus major, a field study Ibis 127: 372-377. Lammens J.J 1967 Growth and reproduction of a tidal flal popula tion of Macoma balihica (L.). Neth. J. Sea Res. 3: 315-382. Lange G. 1968. Uber Nahrung. Nalirungsaulnahme und Vcrdau- ungstrakt mitteleuropaischer Limikolen. Beilr. Vogclk. 13: •{25-334 Laughin R. 1967. Biology of Tipula paludosar, growlh of the larvae in Ihe field. Enl. exp & appl. 10: 52-68. Leeuw JJ. de & M.R. van Eerden 1992. Si/e selection in diving lulled Ducks Asthya fuligula explained by differential han dling of small and large mussels Oreissena polsmorpha. Ardea 80: 353-362
l-cndrem D.W. 1984. Flocking, feeding and predaiion risk: absolute and instantaneous feeding rates. Anim. Behav. 32: 298 299 l.enteren J.C. van & K. Bakker 1976. Functional responses in invertebrates Neth J Zool 26:567-572. I c..p..ui Ml.. C Swennen .c. I. I M. dc Bruijn 1989. Experiments on seleciion on feeding site and lood Size in Ov stercatchers. Haematopus osnalegus. oi different social stains Neth J Sea Re- 23: 333-341. I evinlon J.S. 1971. Control ol lellinacean (MoUusca: Be. feeding behavior by predation. Limnol. Ocean. 16: 660-662. I.evinum J.s. 1991. Variabk : :.ioi III three species of na i Bivalvia. I'elhnccc.i i .is a response lo water flow and sediment transport. Mar. Biol. 110: 375-383. Lima S.L. 1988. Initiation and termination of daily feeding in Dark- eyed Jiincos. influences ol predaiion risk and energy reserve-. Oikos53:3-ll. Lima S.L. & TJ. Valone 1986. Influence of predation risk on diet se lection: a simple example in Ihe Grey Squirrel. Anim. Behav. 34: 536-544 Lin J. & A.H. Hines 1994. Effects of suspended food availability on the feeding mode and burial depth of the Baltic Clam. Macoma balihica. Oikos 69: 28-36. Landers A.M. 1985. Een inleidend onderzoek naar de relatie tussen de fouragerende Scholeksters (Haeinaiopus oslralegus) en de getijdecyclus en hei aanbod van Mossden. Internal student re port 01.85 Rijkswaieisia.il Deitadienst. Middelburg. Lindsay J.A. & N.B Savage 1978. Northern New England's threat ened Soli-shell Clam populations. Environmental Manage ment 2: 443-452. Linke O. 1939. Die Biota des Jadebusenwaltes. Helgolander Meerc- sunters. 1:201-348. Little C. & W, Nix 1976. The burrowing and floating behaviour of the gastropod Hydrobia ulvae. Estuar. csil. mar. Sci.: 4: 537- 544. I uc.is J.R. 198.3. The role of foraging time constraints and variable prey encounter in optimal diet choice. Am. Nal. 122: 191-209. Maagaard L. & K.T Jensen 1994. Prey si/e selection, iniake rate and distnbulion of staging Oystercatchers (Haematopus oslrale- pis) feeding on an intertidal mussel bed [Mytilus edulis). Ophelia Suppl. 6: 201-215. MacAnhur RH. & E.R. Pianka 1966. On optimal use of a palchy en- vironment Am Nai. 100: 603-609. Madsen P.B. & K. Jensen 1987 Population dynamics ol Macoma balthica in Ihe Danish Wadden Sea in an organically enriched area- Ophelia 27: 197-208. Mao P. le 1986. Feeding relationships between Ihe benthic infauna and Ihe dominani benthic fish of ihe Ranee esiuary i France i. J mar. biol. Ass. U.K. 66: 391-401. Masman D. & M. Klaassen 1987. Energy expenditure during free fligbl in trained and free-living Eurasian Kestrels (Falco tin- nunculusi. Auk 104:603-616 REFERENCES 379 Mathivat-Lallicr M.H. & C. Ca/aux 1991 l.ite-hisiory ol Nephtys hombergii in Arcachon Bay. Estuar. Csti Shell Sci. 32: 1-9. M.in'uicsen G.C. I960. Intertidal /onalion in populations ol WjM arenaria. Limnol. Oceanogr. 5: 381-388. Maynes E. & D.I.D. Howie 1985. Biochemical composition and sediment temperature in relation to the reproductive cycle in ihe Lugworm \reiiieola manna. Nelh J. Sea Res. 19: 111-118. McCleery RH 1978. Optimal behaviour sequences and decision making. In: J.R. Krebs & N.B. Davies (eds. I Behavioural ecol ogy: an evolutionary approach: 377-410. Blackwell. Oxford. McGrorty. S..R/I. Clarke, I J Reading & J.D. Goss .('usiard 1990 Population dynamics of the mussel Msiilus edulis density changes and regulation of ihe population in the Exe esiuary. Devon. Mar. Ecol. Prog. Ser. 67: 157-169. McNamara J.M. & A.I. Houston 1986. The common currency for behavioral decisions. Am. Nai. 127: 358-378. Meadows PS. 1964. Experiments on substrate selection by Corophium volululor il'all.isi: deplh seleciion and population dcn-ity 1 Exp. Biol. 41: 677-687. Meadows P.S. & A.A. Ruagh 1981. Temperature preferences and ac- livtiv of Corophium volululor (Pallas) in a new choice appara tus. Sar-ia 66: 67-72. Meiningcr P.L.. A-M. Blomert & E.C.L. Marteijn 1991 Watervo geisterltc in het Dcllagebied. /W-Nederland. gedurende de drie koude winters van 1985, 1986 en 1987. Limosa 64: 89- 11)2. Meire P.M. 1996a. Using optimal foraging theory to determine the density of Mussels Msiilus edulis that can be harvested by hammering OystewalchC""* Haematopus Ostralegus, Ardea 84 in press. Meire P.M. 1996b. Feeding behaviour of Oystercatchers Haemato pus ostralegus during a period of tidal manipulations. Ardea 84: in press Meire P.M. 1996c. I)isinhuii..n ol Oystercalchers Haeinaiopus OS- iralegus ovei a tidal flat in relation to their main prey species, Cockles Ceraslnderma edule and Mussels Msiilus edulis: did ll change after a substanlial habitat loss' Ardea 84 in press. Meire P.M. & A. Ervynck 1986. Are Oysiercatchcis (Haematopus Ostralegus) selecting Ihe mosl profitable Mussels Msiilus edulis)' Anim. Behav. 34: 1427-141-" Mcire P.M.. H. Schekkennan & PL. Meiningcr 1994. Consumption ot benthic invertebrates by waterbirds in ihe Oostersclielde e.luary. SW Nclhcrlands. Hydrobiologia 282/283: 525- 546. Merck T. 1983. Nahrungsokologie des Austernlischers, Haemato pus ostralegus (Avcs. Charadriiformes. Haematopidae): Frei- landsumersuchung zum Beuteerwerb das Wait hesuchender Vogel Diplomarbeil Lniversity ol ( Knabruck. Osnabmck Melcalfe N.B. 1985. Prey detection by intenidally feeding I_apwing. Z. Tierpsychol. 67:45-57. Melcalfe N.B. & R.W. Fumess 1984. Changing priorities: the effect
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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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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 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: llolmann II. & H. Huerschelmann 196
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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