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waders and their estuarine food supplies - Vlaams Instituut voor de ...

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2 3 4 5<br />

probing <strong>de</strong>pth (cm)<br />

Fig. 7. Relationship between probing tunc <strong>and</strong> the <strong>de</strong>pth of a probe.<br />

The values shewn (means ±954 C.L.: sample sizes indicated) are<br />

based on film analysis (resolution l;l(.siol searching behaviour U a<br />

prey <strong>de</strong>nsity of 0 in ": Im probing time) = 0.39 (probing <strong>de</strong>pth)- 2.49<br />

(n = 7. r = 0.97. p < 0.0051.<br />

1 2 3 4 5 6<br />

prey <strong>de</strong>pth (crn)<br />

Fig. 8. Predicted encounter rate (X) per <strong>de</strong>pth class, for three different<br />

prey <strong>de</strong>nsities iprey nr*).<br />

the selected size class, we assume that the bird has to<br />

probe /+1 cm to locate a prey in <strong>de</strong>pth class /. P-loct <strong>de</strong>pends<br />

on D. only, because s is constant for all <strong>de</strong>pth<br />

classes. Dt is known, so Eq (5) can be solved, for t has<br />

been measured using a film analyser (Fig. 7). The logarithmic<br />

increase of probing time with probing <strong>de</strong>pth<br />

OPTIMAL FORAGING AND THE FUNCTIONAL RESPONSE<br />

146<br />

100 200 300<br />

prey <strong>de</strong>nsity (n-nr 2 )<br />

400 500<br />

Ug. 9. Observed mean encounter r.ue I#I t 959 CL. The while<br />

lines represent values of >. ranging from no <strong>de</strong>pth selection (upper<br />

line) to selection ol <strong>de</strong>pth class Ocm only (lower line). The solid linegives<br />

Ihe predicted X for the bird selecting the optimal <strong>de</strong>plh classes<br />

as calculated in Fig. 11.<br />

<strong>de</strong>termines Ihc way the predicted encounter rale <strong>de</strong>creased<br />

for prey which were buried at greater <strong>de</strong>pth<br />

(Fig. 8).<br />

Observed encounter rate<br />

We never saw the Oystercatcher reject a prey it had<br />

discovered ('negalive h<strong>and</strong>ling time"). Thai is why we<br />

used ihe observed searching lime per prey taken to calculate<br />

the mean encounter rate. However, analysis of a<br />

high speed film showed that a prey can be located <strong>and</strong><br />

refused too quickly to be noticed by an observer. We<br />

do not know how often we missed this kind of quick<br />

encounter, bul ii is certain that the observed encounter<br />

rate is an un<strong>de</strong>restimation of me actual rale. Comparison<br />

of the predicted encounter rate for a r<strong>and</strong>om<br />

searching predator <strong>and</strong> the observed rate (Fig. 9) reveals<br />

that at low prey <strong>de</strong>nsities ihe Oystercatcher took<br />

all prey it found, whereas at high <strong>de</strong>nsities less prey<br />

than expected were taken, perhaps because the observer<br />

did not see encounters where prey were rejected.<br />

Another explanation for the lower-than-expected<br />

rate at high prey <strong>de</strong>nsities was suggested by the film

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