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of pre-migralory fattening on the trade-off between foraging and vigilance Behav. Ecol. Soeiobiol. 15:203-206, Mciiam C. 1979. Seasonal changes in populations of Nereis diversi color OV. Milllei from the Severn esiuary. U.K. In: E Nay lor & R.G. Harlnall icds. i Cyclic phenomena in marine planis and animals: 123-130Pergamon Press. New Vork. Moller P. I9S5. Production and abundance ol juvenile Nereis diver sicolor. and oogenic cycle ol adtills in shallow waler- of west- em Sweden. J. mar. biol. Ass. U.K. 65: 603-616. Moller P. & R. Rosenberg 1982. Production and abundance ol Ihe amplnpod Corophium voluiuioi on Ihe west coast of Sweden. Neth. J. Sea Res. 16: 127-140. Moller P. & R. Rosenberg 1983. Recruitment, abundance and pm duciion of Mya armaria and Cardium edule in marine shallow waters, western Sweden. Ophelia 27 Morgan M.J. 1988. The influence ol hunger, shoal si/e and preilaioi presence on loraging in Bluntnosc Minnows. Anim. Behav. 36: 13I7-I322 Mosher J.L 1972. The responses of Macoma balthica (Bh/alvia) to vibrations. Prix malac. Soc. Lond. 40: 125-131. Mourilsen K.N. & K.T. Jensen 1992. Choice ol inicmhabitat in tactile loraging Calidris ulpintc. the importance of sediment penetrability Mai I.e..I Prog Ser. 85: 1-8. Miinch-Pelersen S. 197.1. An investigation of a population ol the Soli Clam (Mya arenaria L.) in a Danish estuary. Medd. Damn. Fisk. -og Havunders.. N.S. 7: 47-73 Murton R.K. 1971 The significance ol a specific search image in Ihe feeding behaviour of the Wood-pigeon. Behaviour 40: 10-42. Murion R.K.. A.J. Isaacson & N.J. Westwood 1971. The signifi cance of gregarious feeding behaviour and adrenal siress in a population oi Wood-pigeons iColiimha palumbus), ). Zool. I ond. 165:53-84. Muus B.J. 1967. The fauna of Danish esiuanes and lagoons. Distri bution and ecology of dominating species in the shallow reaches of the mesohaline zone. Medd. Danm. Fisk. -og Havunders. 5: 1-316. Myers J.P.. S.L. Williams & F.A. Pilelka 1980. An experimental analysis of prey availability for Sanderlings (Aves: Scolopaci- daei feeding on sandy beach crustaceans. Can. J. Zool. 58: 1564-1574. Napolitano G.E. & R.G. Ackman 1989. Lipids and hydrocarbon- in COrophtum volululor from Mimas Basin. Nova Scotia. Mar. Biol. 100:333-338. Nehls G. 1992. Food selection of Knol- Calidris canutus in ihe Wad den Sea: ihe importance ol seasonal and annual variation ol lood availability. Unpublished report. NehI- 0, 1995. Siraiegien der F.rnahrung unci ihre Bedeulung fllr En- ergiehauhall und Okologie der Eiderente (Somateria mollis- suna) (I... 17581 Berichte Forsch. u. Technologie/enirum Westkusied I'mv.Kiel 10: 1-147. Newell CR. & H. Hidu 1982. The effects of sediment type on REFERENCES 380 growlh rate and shell allometry m the Soli shelled Clam t/w; arenaria) J exp.mar.Biol.Ecol.65:285-295. Newell R.C. 1979. Biology of intertidal animals. Marine Ecological Surveys. I aveishaui. Newell R.l.E. & B.L. Bayne 1980 Seasonal changes in Hie physiol ogy. rcprodiicnvc condition and carbohydrate contenl ol ihe Cockle Cardium i= Cerastoderma) edule (Bivalvia: Cardi- idae) Mai. Biol. 56: 11-19. Nie N.H.. CH. Hull. J.G. Jenkins. K. Sleinbrenncr & D.H. Bern 1975. SPSS: statistical package lor Ihe social sciences. Mc Graw-Hill. New York Norton-Griftilhs M. 1967. Some ecological aspccis ol the feeding behaviour of ihe Oysieicalchei Haematopus ostralegus on Ihe Edible Mussel Mytilus edulis, Ibis 109 -112-424. NoruSis M.J. 1988. SPSS/PC+ V2.0 Base manual SPSS Inc.. Chicago. Norusis M.J. 1990. SPSS/PC-f Statistics V4.0, SPSS Inc.. Chicago. Noiusis. M.J. 1993. SPSS for Windows. Base System User's Guide Release 6. SPSS Inc.. Chicago. Ohert B. & H. Michachs 1991. History and ecology ol ihe mussel beds (Msiilus edulis L. i in the catchment area of a Wadden Sea tidal inlet In: M Elliott M. & J-P. Ducrotoy leds.i Esiuanes and coasts: spatial and temporal intercompansons: 185-194. Olsen & Olsen. Frcdenshorg. Olive P.J.W. & RR. Garwood 1981. Gametogenic cycle and popula tion siruciure of Nereis (HediSU) diversicolor and Nereis (Nereis) ,iclegica from north-east England. J. mar. biol Ass U.K. 61: 193-213. Olive P.J.W.. M.G. Bentley. N.H. Wrighl & PJ. Morgan 1985. Reproductive energetics, endocrinology and population dy namics of Vephlys i aei a and N hombergl. Mar. Biol. 88: 235- 246. Orth R.J. 1975. Destruction of eelgrass. Zoslera marina, by the Cownose Ray, Kluiiopieru bonasus, in Ihe Chesapeake Bay. Chesapeake Sci. 16:205-208. Oyenekan J.A. 1986. Population dynamics and secondary poduction in an estuarine population of Nephtys hombergil I Polychaela: Nephtyidaei. Mar Biol 93:217-223. Paine R.T. 1966. Endoihermy in bomb calorimetry. Limnol. Ocean. II: 126-129, Pen son W.H.. D.L. WoodrulT. P.C. Sugarman & B.L. Olla 1981 Ef fects of oiled sediment on predation on the Littlcncck Clam. Pniioiliacu staminea, by the Dungeness Crab, Cancer magis- ter. Estuar. Csil. Shelf Sci. 13: 445-454. Pehrsson O. 1976. Food and feeding grounds of the gold. Unci phalli clungula tL.I on Ihe Swedish wesl coasi. Onus Scand. 7:91-112. Pekkarinen M. 1983. Seasonal changes in condition and biochemi cal constituents in the soft pan ot Macoma balihica (Lamelli- branchiata) in the Tvarminne brackish waler area (Baltic Sea). Ann. Zool. l-enn. 20:311-322.
Pekkarinen M. 1984. Regeneration of the inhalant siphon and siphonal sense organs ol brackish water (Baltic Seal Macoma balthil a (Lamellibiaiichiala. Tellinaceal Ann. Zool. Fcnn. 21. 29-40 Pennycuick C.J 1989. Bird flight performance: a practical calcula tion manual. Oxford University Press. Oxford. Peterson CJi & M.L. Quammen 1982. Siphon nipping us impor tance to small fishes and its impact on growth ol ihe bivalve Prololhaca slaminea (Conrad). J exp. mar Biol Ecol. 63: 249-268 Picnkowski M.W. 1977. Difliences In babital iei|uiieinenis and dis tribution pattern of plovers and sandpipers as investigated by studies ol leeduig behaviour Verb om ties. Bayern 23-105 174. Picnkowski M.W. 1983a. The effects of environmental conditions on feeding rates and prey seleciion of shore plovers. Ornis Scand. 14:227-238. Pienkowski M.W. 198.3b. Surface activity of some intertidal inverte brates in relation to lemperaliire and the loraging behaviour of their shorebird predators. Mar. Ecol. Prog. Ser. II: 141-150. Piersma T 1987. Production by intertidal benthic animals and limits lo their predation by shorehirds: a heuristic model. Mar Bcol Prog. Ser. 38: 187-196. Piersma T 1989. Seasonal changes in the food requirements of Knots. Annual report 1988: 22-23 Netherlands Institute lor Sea Research, Texel. Piersma T. 1996. Energetic con-nauiis on the non-breeding dislnhu- lion of coastal shorehirds. Inleniational Wader Sludies s' 122- 135. Piersma T. L. Zwarts ,Si J II Bruggemann 1990. Behavioural as- pects of the departure ol waders before long-flight distances: flocking, vocah/alions. flight palhs and diurnal timing. Ardea 78: 157-184. Piersma I.. R. Drent & P. Wiersma 1991. Temperate versus tropical wintering in Ihc world's northernmost bleeder, the Knol. metabolic scope and resource levels restrict subspecies op tions. Acta XX Congr. intern. Ornith.. Christchurch: 761-772. Piersma T. A. Koolhaas & A. Dekinga 1993a. Interactions between stomach Structure and diet choice in shorehirds. Auk 110: 552- 564. Piersma T, R. Hockstra. A. Dekinga. A. Koolhaas. P. Wolf. P. Battle] & P. Wiersma 1993b. Scale and intensity of intertidal habitat use by Knots Calidris. anutus in the western Wadden Sea in re lation U> food, friends and foes. Neth. J. Sea Res. 31: 331 -357. Piersma I . Y. Verkuil & I. Tulp 1994 Resources for long-distance migration ol Knots Calidris caniiius islandica and C. c. canii ius: how hroad is the temporal exploitation window of benthic prey in the western and eastern Wadden Sea'' Oikos 71: 393- 407. Pier-ma T. N. Cadee & S. Daan 1995. Seasonality in basal metabolic rale and thermal conductance in a long-distance mi REFERENCES 381 grant shorebird. the knot (Calidris canutus). J. Comp. Physiol. B 165; '7-45. Piersma I.. L. Bruin/eel. R. Drent. M. Kersten. J.v.d. Meei. I 1 Wiersma, 1996. Variability in basal metabolic rales ol a long distance migrant shorebird (Red Knol. Calidris canutus) lellecl shllls in ol-ail si/es Physiol /oo| Is'l |.|| J[7 Pietcrs H.. J.H. Kluylmans, W. Zurburg* D.I. Zandee 1979. The in fluence of seasonal changes on energy iiieiabolism in Msiilus edulis (L.i. I. Growlh rate and biochemical coiiiposiiinn in re lation to environmental parainelci- and spawning. In 1 lor& R.G. Hamuli (eds.) C ychc phenomena in marine plains and animals: 285-293. Pergamon Press. New York. Pietcrs H.. J.H. Kluylmans & D.I. Zandee 198(1 1 issue composition and reproduction ol Msiilus edulis in relation to lood availabil ity. Neth. J. Sea Res. 14: 349-361. Pihl L. 19X2. Food iniake of young Cod and Rounder in a shallow bay on ihe Swedish wesl coast Neth J. Sea Res. 15: 419-432. Pihl L. & R. Rosenberg 1984. Food seleciion and consuinpiion of Ihe shrimp Crangon cniiigon in some shallow marine areas in •westernSweden. Mai Ecol Progr. Ser. 15: 159-168. Pitcher T.J.. S.H. Lang & J.A. Turner 1988. A risk-balancing trade off between loraging rewards and predaiion hazard in a shoal ing fish. Behav. Ecol. Sociobiol. 22: 225 Prater AJ 1972. Die ecology of Moreeambe Bay 111. The food and feeding habils of Knot (Calidris caniiius L.) in Moreeambe Bay. J. appl. Eel. 9: 179-194. Prater A.J. 1981. Esiu.uy buds of Britain and Ireland. Poyser. Cal- lon. Prokosch P. 1988. Das Schlcswig-Holsteinische Watlenmeer als I nihiahrs-.Aulenhallsgebiel Arklischer Walvogd-Populatio- nen am Beispiel von Kiebiizregenpleiler il'luvialis iqualarola, L. 1758). Knutl (Calidris caniiius. L. 1758) und Puhlschnepne (Limosa lap/miuca. L. 1758). Cora*C 12: 273-442. I'll 11 i.in l H.R. 1980. Do Chipping Spanow s forage optimally' 1 Aidca 68: 75-82. Ralcliffe PJ.. N.V. Jones & N.J. Wallets 1981. The survival of Ma- coina balthica (Ll in mobile sediments. In: N.V. Jones & W.J. Wolff (eds. I Feeding and survival strategies ol estuarine organ isms: 91-108 Plenum Press. New York. Reading CJ. & S. McGrorty 1978. Seasonal variations in the bury ing depth of Macoma balthica (L. land iis accessibility to wad ing birds. Estuar. csil. mar. Sci (v 135-144 Rechten C. M. Avery & A. Stevens 1983. Optimal prey selection: why do Great fits sh. m partial preferences? Anim. Behav. 31: 576-584. Reddingius H . A.J Schilstru & G. Thomas 1983. The grid method in estimating the path lengih ol a moving animal. J. Anim. Ecol. 52: 199-208. Rcid G.B. & A. Reid 1969. Feeding processes ol tnemtvrs of the genus Macoma i MoUusca Bivalvial Can. J Zool. 47:649-657. Reise K. 1977. Predalor exclusion experiments in an intertidal mud
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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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VERSATILITY OF CURLEWS FEEDING ON N
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Page 331 and 332: PREY DEPLETION BY OYSTERCATCHER AND
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Page 343 and 344: Voorgeschiedenis In 1960 verscheen
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Page 349 and 350: kunnen opzuigen. of diep leven en z
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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 and 374: llolmann II. & H. Huerschelmann 196
Page 375: l-cndrem D.W. 1984. Flocking, feedi
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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