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I ustart 1.1). A I) Wtat, R.T. Clarke. R.W.G. Caldow. S.E.A. le V. dil Durell I996d. The cany ing capacity ol coastal habitats for Oystercatchers. In: J.D. Gos--Custard (ed.) The Oyster- calcher: from individuals lo populations: 327-351. Oxford University Press. Oxford Goss-Custard J.D.. S.E.A. le V. dit Durell. R.T. Clarke. AJ. Bein- tema. R.W.G. Caldow. PI . Meuiinger&CJ. Smit I99(K- Pop ulation dynamics: predicting the consequences ol habitat change at the continental scale. In: J.D. Goss-Cuslard (ed I 1 he Oystercatcher from individuals lo populations: 352-383 Ov i..nl i invcrsiiv Press,Oxford. Goullelqucr P. & M. Wolowicz 1989. The shell of Cardium edule, Cardium glaucum and Ruditapes philippinariim. organic con tent, composition and energy value, as determined by different methods. J. mar. hiol. Ass. I K 69: 56 (li.iiio G.W.. L.H. Thomas & CL. Graiio 1984 Some aspects of the foraging ecology of migrant juvenile sandpipers in the outer Bay ..t Fundy. Can. J. Zool. 62: 1889-1892. Green J. 1967. Activities of the siphons ot Scrobicularia plana (da Costa) Proc. malac Soc. Lond. 37:339-341. Green J. 1968. The biology ol estuarine animals. Sulgwick & lack son. London. Green R. 1978. Factors affecting ihe did of farmland Skylarks Manila arvensis. J. Anim. Ecol. 47: 913-928. GrubbT.C. & L. Greenwald 1982. Sparrows and a bnisfipile: Forag ing responses to different combinations of predation risk and energy cost. Anim. Behav. 30: 637-640. Habekolle B. 1987. Scholeksters en Slijkgapers. Student report /,» L.gual 1 aboratory. University of Groningen. Groningen. Haddon M.. R.G. Wear * II \ Packer 1987. Depth and density ol burial by ihe bivalve Paphies ventricosa as refuges from preda iion by the crab Uvalipes catharsis. Mar. Ecol 94 Hancock D.A. 1967. Growlh and mesh selection in the Edible Cockle (Cardium edule L.I. J. appl. Ecol. 4: 137-157. Hancock D.A. & A. Franklin 1972. Seasonal changes in ihe condi lion ol the Edible Cockle (Cardium edule I..). J. appl. Ecol 9: 567-579. Hancock D.A. & A.E. Urquhart 1964. Mortalities of Edible Cockles (Cardium edule Li during Ihe severe winter of 1962-63. J. Anim. Ecol. 33: 176-178. Hancock D.A A Al Urquhart 1965 The determination ol natural mortality and its causes in an exploited population of Cockles (Cardium edule L.I Fish Invest Minisi Agric Fish Food Ser. 1124: 141). Harris M.P. 1965. The lood of some Lams gulls. Ibis 107: 43-53. Harley MB. 1950. Occurrence of a filter feeding mechanism in the Polvehacle Nereis diversicolor. Nature, l.ond. 165: 735. Harley M.B. 1953. The feeding habits of Nereis diversicolor, Br J Anim. Behav. I: 88. Harvey M. & B. Vincenl 1989. Spatial and temporal variations ol the re-production cycle and energy allocation of the bivalve Ma- REFERENCES 376 mum balthk a (Li on a tidal flat. J. exp. mar. Biol. Ecol. 129: 199-217. Harvey M. & B. Vincenl 1990. Density, size distribution, energy al location and seasonal variation in shell and sofl tissue growth at two tidal lev els oi a Macoma bahhica (L.) population. J. exp. mar. Biol. Ecol. 142: 151-168. Ha-sell M.K J II. Law ton & J.R. Beddington 1976. The components of arthropod predation I The prey death rale. J. Anim. Ecol. 45: 135-164 Hassell M.P. J.H. Lawton & J.R. Beddinglon 1977. Sigmoid func tional responses hy invertebrate predators and parasiloids. J. Anim. Ecol. 46: 249-262. Hawkins CM. 1985. Population carbon budgets and the importance of the amphipod Corophium voliiialor in ihe carbon transfer on a Cumberland Basin mudflat, upper Bay of Fundy. Canada Neth. J. Sea Res, 19: 165-176. I lay nes DL. & P. Sisojevic 1966. Predatory behav iour of PhUodro- inns rufus vValkenaer (Aranea: Thomisidael Can. Em. 98: 113-133. Ileezik V.M van. A .EC. Gerritsen & C. Swennen 1983. The influ ence of cheinoreceplion on the loraging helutvioui ..I two species ol sandpiper Calidris alba and Calidris ulpina. Neth. J. Sea Res. 17:47-56. Ilcgner R.E. 1985. Dominance and anti-predator behaviour in Blue Tils (Panis caeruleus). Anim. Behav. 33: 762-768. Heller R. & M. Milinski 1979. Optimal loraging of sticklebacks on swarming prey Anim Behav 27: 1127-1141. Henderson S.M. & CA. Richardson 1994. A comparison of ihc age. grow ih rale and burrowing behaviour of razor dams, Ensis till qua and E ensis. J mar. biol. Ass. U.K. 74: 939-954. Hepplcsion P.B. 1971. The feeding ecology of Oystercatchers (Haematopus oslralegus L.i in winter in northern Scotland. J Anim. Ecol. 40: 651-672. Hibbert CJ. 1976. Biomass and production of a bivalve community on an intertidal mud-flat. J. exp. mar. Biol Ic.l 25: 249 261. Hicklin P.W. & P.C. Smith 1984. Selection ol loraging sites and in vertebrates prey hy migrant Semipalmaied Sandpipers Calidris pusilhi (Pallas) in Minas Bay. Bay of Fundy. Can. J. Zool, 62: 2201-2210, Hodgson A.N. 1981. 'Die blood volume of ,V< robiculanaplana, and an estimation of blood loss after siphonal wounding. Mar. Behav. Physiol. 8:21-33. Hodgson A.N. 1982a. Studies on wound healing, and an estimation of ihe rale ol regeneration, ol ihe siphon of Scrobicularia plana (da Costal. J. exp. mar. Biol. Ecol 62: 117-128. N. 1982b. Some behavioural am responses of Scrobicularia plana iBivalvia: Tellinaceal lo siphonal ".mind ing J. Moll. Stud. 48: 87-94 Hodgson A.N. & E.R. Trueman 1981. The siphon- ol Scrobicularia plana iBivalvia: Tellinaceal. Observations on movement ami extension. J. Zool. Lond. 194: 445-459.
llolmann II. & H. Huerschelmann 1969. Naluiingsuntctsuchungen bei Limikolen durch MagemnhalLsanalysen. Corax 3: 7-22. Holbrook SJ. & R.J. Schmitl 1988, Effects ol predaiion risk on for aging behai ion mechanisms altering patch choice. J. exp. mar. Biol. Ecol. 121: 151-163. Holland A P..N.K Mountfonl, M.H Hiegel. K.R. KaumeyerA J.A. Mihursky I9S0. Influence of predaiion on inlaunal abundance in upper Chesapeake Bay. USA. Mar. Biol. 57: 221-235. Holling C.S. 1959. Some cli.uuclciisiicsol simple types of predaiion and parasitism. Can. Enl. 91: 385-398. Holling C.S. 1966. The funcin.ii.il response of invertebrate predators to prey density. Mem. Enlomol. Soc. Can. 48: I -86. Holihuijzen Y.A. 1979. Hei voedsel van de /wane Ruiter Tringa erythropus in de Dollard. Limosa s2 22-33. Hosper U.G. 1978. Fouragccrstralegie en voedselopname van Scholeksiers (Haematopus ostralegus) in bet binnenland Stu dent report Zoological l.aboiatoiy. University ol Groningen. Groningen. Hughes R.N. 1969. A study of feeding in Scrobicularia plana. J mar. biol. Ass. U.K. 49: 805-823 Hughes R.N. 1970a. Population dynamics of the bivalve .Si robicu- UH ni plana (da Costa) on an intertidal mud-flat in North Wales. J. Anim. Ecol. 39: 333-356. Hughes R.N. 1970b. An energy budget for a tidal-flal population of Ihe bivalve Scrobicularia plana Ida Costal. J. Anim. Ecol. 39: 357-381. Hughes R.N. 1972. Reproduction ol Scrobicularia plana da Costa (Pelecypoda: Semelidaei in North Wales. Veliger 14: 77-81. Hughes R.N. 1979. Optimal diets under the energy maximization premise: Ihe effects of recognition time and learning. Am. Nat 113: 209-221. Hughes R.N. 1980. Optimal loraging theory in the marine context Oceanogr. Mar. Biol. Ann Rev 18: 423-481. HughesT.G 1973. Deposii feeding in Abra tenuis ( Bivalvia. pp. Tel- linacea). J. Zool. Lond. 171: 499-512. Hulscher J B. 1964. Scholeksier en Lamellihraiichialen in de Wad denzee. Levende Natuur 67: 80-85 Hulscher J.B. 1973. Burying-depth and trematodc infection in Macoma balihica. Neth. J. Sea Res. 6: 141 -156 Hulscher J.B 1974. An experimental study of the food iniake ol the Oystercatcher Haematopus ostralegus L. in captivity during the summer. Ardea 62: 156-171. Hulscher J.B. 1976. Localisation of Cockles I Cardium edule L i by ihe Oystercatcher i Haematopu s ostralegus L.) in darkness and daylight. Ardea 64: 292-310. Hulscher J B 1982. The Oystercatcher Haeinaiopus oslralegus as a predator of the bivalve Macoma balihica in the Dutch Wadden Sea. Ardea 70:89-152 Hulscher J.B. 1988. Mossel doodl Scholekster Haematopus ostrale gus. Limosa 61: 42-45, Hulscher J.B. 1990. Survival of Oystercalchers during hard winter REEERENCES 377 weather. Ring 13: 167-172. Hulscher J.B. 1996. Food and feeding behaviour. In: ID. Goss-Cus- lard led.I The Oysiercalcher: from individuals to populations: 7-29 Oxford Lniversitv Press. Oxford. Hulscher J.B. & BJ. Ens 1991. Somatic modifications ..I feeding system structures due to feeding on different foods will phasis on changes m bill shape III Oystercatchers. tcta \\ ( ongr, intern Ornilh . Christchurch: 889-8%. Hulscher J.B. EJ. Bunskoeke. D. Ailing & BJ. Ens 1996. Subtle differences between male and female Oystercatcher Haemato pus OStlldegUS bl locating and handling the bivalve mollusc Ma oiiiii balihica, Ardea K4: in press Hummel II. 1985a Pood intake ol M.iionui balthica l Mollusc.n ill relation K) seasonal changes in its potential food on a tidal flat in Ihe Dutch Wadden Sea. Neth. J Sea Res. 19: 5 Hummel H. 1985b. Fi*xl intake and growlh in Macoma balthica i MoUusca) in the laboratory. Neth. J. Sea Res. 19: 77-83. Hummel H. 1985c. An energy budget of Macoma balihica iMol- luscai population living on a tidal flat in the Dutch Wadden lb J. Sea Res 10 s Jackson MJ. & R. James 1979. The influence of bail digging on Cockle. Cerastoderma edule. populations in North Norfolk. J. appl. Ecol. 16:671-679. Jensen KT. 1985. The presence ol the bivalve (. erastoderma edule affects migration, survival and reproduction of ihe amphipod ( oropliium votutator. Mar. Ecol. Prog. Ser. 25: 269-277. Jensen KT. & J.N. Jensen 1985. The importance ol some epibenihic predators on the density of juvenile benthic maerofauna in the Danish Wadden Sea. J. exp. mar. Biol. Ecol. 89 157-174. Jensen KT. & E.D. Krisiensen 1990. A field expenmenl on compe tition between Corophium volutalor (Pallasl and Corophium armarium Crawford (Crustacea: Ampinpodai: effects oo sur vival, reproduction and recruitment. J. exp. mar. Biol 137: 1-24. Johnson C. 1985 Patterns of seasonal weight variation in waders on ihe Wash. Ring. & Migr. 6: 19-32. Johnson R.G. 1965. Temperature variation in the inlaunal environ- nieni of :i sand Hal. Limnol. Oceanogr 1(1: 114-120. Jones A.M. 1979. Structure and growth of a high-level population ot Cerastoderma edule il.amellibranchiatai. J. mar. biol. Ass U.K. 59:277-287. Jones D. & J. Clare 1977. Annual and long-term fluctuations in the abundance of fish species inhabiting an intertidal mussel bed in Moreeambe Bay. Lancashire Zool. J. Limn. Six:. 60: 117-172. Jones N.V. & C. Park 1991. A population of Macoma balilu studied over 16 years in the Humber estuary In M Elliott & J- P. Ducrotoy (eds l Estuaries and coasts spatial and temporal iiiiercoiiipans.in. 153-157. Olsen & Olsen. Fredcnshorg Jonge V.N. de & K. Essink 1991. Long-term changes in nutnenl loads and primary and secondary producers in the Dutch Wad den Sea. In: M. Elliott & J-P. Ducrotoy (eds.) Estnaries MIS\
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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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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: huch der Vogel Milteleuropas, Band
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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