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Esselink P & I. /warts 1989. Seasonal trend in burrow deplh and lidal variation in feeding activity of Nereis diversicolor. Mar. Ecol. Prog. Ser. 56: 243-254. Esselink P.. J van Belkum & K. Essink 1989. The effect of organic pollution on local distribution of Nereis diversii olor and Corophium vohaator. Neth. J. Sea Res. 23: 323-332. Essink K. 1978. The effects ol pollution hy organic waste on macro- fauna in the eastern Duich Wadden Sea. Nelh. Inst Sea Res. Publ. Ser. 1-1978: 1-135. Essink K. & A.H. Bos 1985 Growlh of three bivalve molluscs trans planted along the axisol the Ems estuary. Neth. J. Sea Res. 19 45-51. Essink K.. H.L. Kiev!. W Vi.scr& P.Tydcman 1985. Population dy namics of the Ragworm Nereis diversicolor in the Dollard 11 ins estuary i under changing conditions of stress hy organic pollution. In: J.S. Gray & M.E. Christiansen teds.) Marine biology of polar regions and effects ol stress on marine organ isms: 585-600. John Wiley. Chiche.ler. Essink K.. J J. Beukema. J. Coosen. J.A. Craeymeersch. J-P. Ducro toy. H. Michaelis .V |j. Robineau 1991. Population dynamics ol the bivalve mollusc Scrobicularia plana da Costa: compar isons in lime and space. In: M. Elliott & J-P. Ducrotoy icds.i Estuaries and coasts: spatial and temporal intcrcomparisons: 167-172. Olsen & Olsen. Predensborg. l-.uiernioser A. 1961. Schlagen Beizlalken bevor/ugl kranke Krahen ' Vogdwell 82: 101-104. Evans A. 1987. Relatively availability ol the prey of wading birds hy day and by night. Mar. Ecol. Progr. Ser. 37: 103-107. Ivans PR. 1979. Adaptations shown by foraging shorehirds to cycli cal variations in the activity and availability of their intertidal invertebrate prey. In: E. Naylor & R.G. Hartnall (eds.) Cyclic phenomena in marine plains and animals: 357-366. Pergamon Press, New York. Evans PR 1988 Predaiion of intertidal fauna hy shorehirds in rela tion lo time of the day. tide and year. ln:G. Chelazzi &M. Van illin (eds.i Behavioral adaptation io inienidal life: 65-78. Plenum Press. New York Evans P.R.. D.M. Herdson. D.J. Knight & M.W. Pienkovvski 1979 Short-term effects of reclamation of pan ol Seal sands. Teesmouth. on wintering waders and Shelduck Oecologia (Beit) 41: 183-206 Evans S.M A: I Rogers 1978. Organization of lubiculous behaviour in ihe polychaelc Nereis diversii olor (OJ?. Miiller). Mar. Be hav. Physiol. 5: 271-288. Everards K. 1973. IX* groei van Msiilus edulis L. als functie van de onderdompdingsiijd. NIOZ report 1973-2 Netherlands Insti tute for Sea Research. Texel Farke H. & E.M. Berghuis 1979. Spawning, larval development and migration ol Arenicola marina under field conditions in Ihe western Wadden Sea. Neth. J Sea Res. 13: 529-535. Feare CJ., G.M. Dunnet & IJ. Patterson 1974. Ecological studies of REFERENCES 374 ihe Rook K ..mi. Intgilegiis L.) in North-East Scotland: food iniake and feeding behaviour J. appl. Ecol. 11: 867-8%. I isher II. 198.3. Shell weighl as an independent variable in relation to cadmium content of molluscs. Mar. Ecol. Prog. Ser. 12: 59- 75. I uzCibbon CD. & J. H Fanshawe 1989. The condition and age of Thompson's gazelles killed by cheetahs and vi ild dogs. J. Zool. Lond. 218:99-107. Flach E.C. 1992. The influence of lour macrozoobenthic species on the abundance of the amphipod Comphium volutaior on tidal flats in the Wadden Sea. Nelh. J. Sea Res. 29: 379-394 Frelwell SD. 1972 Populations in a seasonal environment. Prince ton I nivei-nv Press, Princeton, New Jersey. GayaniloF.C Jr.. M.Soriano ft D.Pauly 1988. ,\ draft guide to the Compleal EI.EFAN 1(1 ARM Software 2. International Cen ter for Living Aquatic Resources Management. Manila. Gendron R.P. 1986. Searching for cryptic prey: ev ulence for optimal search rales and the formation of search images in Quail. Anim. Behav. 34: 898-912. Gendron R.P. & J.E.R. Staddon 1983. Searching for i ry puc piey the effect ol search rate. Am. Nat. 121: 172-186. Gerritsen A.F.G 1988 General and specific patterns in loraging acts of sandpipers (Calidris). In: A.F.G. Gerritsen Feeding tech niques and the anatomy ol ihe hill in sandpipers {Calidris): 91- 137. Thesis University of Leiden, leiden Gerritsen A.F.G. & A. Mciboom 1986. The role of touch in prey density esiimaiion by Calidris alba. Nelh. J. Zool. 36: 530- 562. Gerritsen A.F.G.. Y.M. van Hcezik & C. Swennen 1983. Chemore- ception in two further Calidris species {('. maritimus and C. canutus) with a comparison of Ihe relative importance of cheinoreception during foraging in Calidris Species, Neth. J. Zool. 33:485-496. Getty T. & H.R. Pulliam 1993. Search and prey detection by forag ing Sparrows. Ecology 74: 734-742. Gilbert M.A. 1973. Growth rale, longevity and maximum size ol Macoma balihica l\..). Biol Bull 145: 119-126. Gilbert M.A. 1977. The behav IOIII and functional morphology of de posit feeding in Macoma balihica (Linne. 1758). in New Eng land. J. Moll. Stud. 43: 18-27. Giller PS. 1980 The control of handling time and its effect on the loraging strategy of a heleroptcran predator, Notonecla. J. Anim. Ecol. 49: 699-712. Goede A.A. 1993. Variation in Ihc energy intake of captive oyster catchers Haeinaiopus oslralegus. Ardea 81: 89-97. flli.np.-r R I n*e 1967 Over de bodemgesteldh.'>i \ in bet wad dengebied Van Zee to Land no. 43. Tjecnk Willink. Zwolle. Gliick E. 1987. An cxperimeni.il study ol feeding, vigilance and predator avoidance in a single bird. Oecologia tBerl.l 71: 268- 272. Glutz von Blotzhcim U.N.. K.M. Bauer & I-. Bezzel 1975. Hand-
huch der Vogel Milteleuropas, Band o Akademische Vcrlags- gesellschafl. Wiesbaden. Goede A.A. 1993. Variation in the energy iniake ot captive Oysier S9 97. Goerke H. 1966. Nahrungsliltration von Nereis diversicolor O.F. MOJIer i.Nereidae. Polychael.n vcioll. Insi Meeiesloisch. Bremerh. 10:49-58. Goerke H. 1971. Die Ernahrungsweisc der Nereis-Ancn (Poly- chaeta. Nereidae) der deiilschen Kiisien Verolt. Insi Meeres- forsch Bremerh. 13: 1-50. Goss-Custard J.D. 1969. The winter feeding ecology of the Red shank Tringa totanus. Ibis 111: 338-356. (loss (usiard J.D. 1970a. The responses of Redshank (Tringa totanus) io spatial variations in the density of their prey. J. Anim. Ecol. 39:91-113. Goss-Custard J.D. 1970b. Feeding dispersion in some overw mienng wading birds In: J.H. Crook (ed.) Social behaviour in birds and mammals: 3-35. Academic Press. London. GoSS-Custaid J.D- 1976. Vanation in the dispersion of Redhank Tringa totanus on llieir winter feeding grounds Ibis 118: 257- 263. Goss-Custard J.D. 1977a. The energelics ol prey seleciion by Red shank. Tringa loiiinus (L.|. in relation lo prey density. J. Anim. Ecol. 46: 1-19. Goss-Custard J.D. 1977b. Optimal foraging and ihe size selection ol worms hy Redshank. Tringa totanus, in Ihe field. Anim. Behav. 25: 10-29. Goss-Cusiard J.D. 1977c. The ecology of ihe Wash III. Densiiy re lated behaviour and the possible effects of a loss of feeding grounds on wading birds (Charadrii). J. appl. Ecol. 14: 721-739. GoSS-CuStard J.D. I977d. Predator responses and prey mortality in the Redshank Tringa totanus il I and a preferred prey Corophium volutator I Pallas). J. Anim. Ecol. 46: 21-36. Custard J.D. 1980. Competition Im food and interference among waders Ardea 68: 31-52. ( ustard J.D. & S.E.A. le V. dil Durell 1983. Individual and age differences in ihe feeding ecology of Oystercatchers Huemato- pusostralegus w mtering on ihe Exe estuary Ibis 125: 155-171. Goss-Custard J.D. & S.E.A le V. dil Durell 1984. Feeding ecology. winter mortality and the population dynamics ol Oysiercatch- ers on the Eve estuary. In: P.R. Evans. J.D. Goss Custard. W.G. Hale (eds.) Coastal waders and wildfowl in winler: 190-208. Cambridge University Press, Cambridge Goss-Cuslard J.D. & S.E.A. le V. dil Durell 1987. Age-related ef- lects in Oystercatchers. Haematopus oslralegus. feeding on Is, Mytilus edulis. I. Foraging efficiency and interfer ence. J. Anim. Is.>! 56 "2 1-5.36.
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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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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 and 368: BreyT. 1989. Der Einfluss physikali
Page 369: latie tol chironoiiiiilen en de wai
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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