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

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1 2 3 4 5 6 7<br />

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

PREY SIZE SELECTION AND INTAKE RATE<br />

350 prey nv 2<br />

0 1 2 3 4 5 6 7<br />

maximal prey <strong>de</strong>pth accepted (cm)<br />

Fig. 12. Experiment toshow why an Ov siercatchei ignored <strong>de</strong>ep, less profitable prey when the intake rale increased due to a high prev <strong>de</strong>nsity<br />

V. Tune nee<strong>de</strong>d to h<strong>and</strong>le Scrobicularia 37-38 mm long which were laken from different <strong>de</strong>pths (s ± SE). B. Time nee<strong>de</strong>d io probe to different<br />

<strong>de</strong>pths (s ± SE). C. Predicted feeding rate (Scrobicularia min ') at two prey <strong>de</strong>nsiiies when an Oystercatcher took prey from the upper I. 2. ...<br />

7 cm of the substrate. Actual observed <strong>de</strong>pth selection is indicated vv ith horizontal bars. As predicted, the bird took prey from all <strong>de</strong>pths when<br />

the prey <strong>de</strong>nsity was low <strong>and</strong> only shallow prey when the prey <strong>de</strong>nsity was high However, the bird did this al a much higher rate than predicted<br />

(explanation given in text). Data of Wanink & Zwarts (Iis s ><br />

nomics of foraging are being calculated.<br />

It has been shown experimentally that a captive<br />

Oystercatcher <strong>de</strong>creased its probing <strong>de</strong>pth from 7 to 3<br />

cm when the <strong>de</strong>nsity of the prev on offer. Scrobicularia<br />

36-37 mm long, increased from 24 to 350 prey nr 1<br />

(Wanink & Zwarts 1985). It took more time to h<strong>and</strong>le<br />

<strong>de</strong>ep-living prey (Fig. I2A). hence prey profitability<br />

<strong>de</strong>creased with <strong>de</strong>pth. Moreover, it also took more time<br />

in the first place to locate a prey at greater <strong>de</strong>pths (Fig.<br />

I2B). The encounter rate, <strong>de</strong>fined in optimal foraging<br />

mo<strong>de</strong>ls as the inverse of the searching time, for prey at<br />

different <strong>de</strong>pths could be calculated from the effective<br />

touch area <strong>and</strong> the probing time at each probing <strong>de</strong>pth.<br />

Since the encounter rate <strong>and</strong> the h<strong>and</strong>ling time were<br />

both known for each <strong>de</strong>pth class, the optimal set of<br />

<strong>de</strong>pth classes, which should be inclu<strong>de</strong>d in the diet to<br />

maximize the intake rate, could be predicted exactly<br />

with a multi-species functional response equation<br />

(Charnov 1976) (Fig. 12C). The <strong>de</strong>pth selection ma<strong>de</strong><br />

166<br />

by the bird was very close to that which was predicted.<br />

The Oystercatcher took prey from all <strong>de</strong>pths when the<br />

<strong>de</strong>nsity was low but rejected the <strong>de</strong>ep, less profitable<br />

prey when prey <strong>de</strong>nsity was high: by doing so, it increased<br />

its intake rate at high prey <strong>de</strong>nsity by eating<br />

only the most profitable, shallow prey. It may be<br />

thought from Fig. 12C that the bird could have done<br />

slightly better by selecting prey only from the upper 1<br />

cm, <strong>and</strong> not from the upper 3 cm. as it actually did.<br />

But. in fact, the Oystercatcher did even better than predicted<br />

by using a second selection criterion, as will be<br />

explained in the next section. The conclusion from this<br />

experiment is that Oystercatchers are in<strong>de</strong>ed able to<br />

vary the rate at which they encounter different prey<br />

types <strong>and</strong>. in doing so, increase <strong>their</strong> intake rate.<br />

Rejection of prey to increase intake rate<br />

The previous section showed that Oystercatchers are<br />

able to adjust <strong>their</strong> searching behaviour in or<strong>de</strong>r to in-

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