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

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IOO<br />

BO<br />

60<br />

40<br />

20<br />

0<br />

PREDICTING SEASONAL AND ANNUAL FLUCTUATIONS IN THE LOCAL EXPLOITION OF DIFFERENT PREY<br />

0 loss ol body weighl<br />

| mass mortality Cod<strong>de</strong>s<br />

• mortally (unknown reason)<br />

| predation Oystercatcher<br />

ii?<br />

S?:A<br />

70 80 90 100 110 120<br />

biomass at 15 August (guv 2 )<br />

Ki 1 *. 18. Loss of biomass by Cockles. Scrobicularia. Macoma. Mya<br />

<strong>and</strong> Mussels between 15 August <strong>and</strong> 15 March of the next year in rehiiion<br />

to the total biomass present at 15 August. The loss of biomass<br />

is due to a <strong>de</strong>crease in Ihe body condilion <strong>and</strong> to mortality, the latter<br />

given separately for Oysiercatchcr predation, mass mortality in<br />

Cockles (see text) <strong>and</strong> other causes. Panel A. gives the absolute loss<br />

of biomass (g m : ). <strong>and</strong> B. the loss as percentage.<br />

when the <strong>food</strong> supply was poor <strong>and</strong> this increased to<br />

17% in winters with a rich <strong>food</strong> supply.<br />

Discussion<br />

Mo<strong>de</strong>l assumptions<br />

In this paper we predicted the feeding behaviour of the<br />

Oystercatcher on the basis of measurements on the<br />

prey. This required several simplifying assumptions.<br />

Before we discuss our results in some <strong>de</strong>tail, it seems<br />

pru<strong>de</strong>nt to investigate some limitations in our approach.<br />

256<br />

Spatial variation In our mo<strong>de</strong>l calculations we ignored<br />

the spatial variation in prey <strong>de</strong>nsity. The samples<br />

of the macrozoobenthos in the Nes area were taken at<br />

73 sites <strong>and</strong> we know from this mat Macoma <strong>and</strong> Scrobicularia<br />

on the rich sites were twice as common as on<br />

the poor sites <strong>and</strong> that the ratio was even three in Cockles<br />

<strong>and</strong> Mya (Zwarts 1988b). In principle, it would<br />

have been possible to repeat all calculations done for<br />

the Nes area as a whole for the 73 sites individually.<br />

This would not have changed the average trends shown<br />

for prey choice <strong>and</strong> intake rate. However, whereas we<br />

predicted that all birds would everywhere have<br />

switched at the same moment from one prey to the<br />

other. Ihe timing would have been different between<br />

sites, so that more gradual changes would be expected<br />

for the Nes area as a whole.<br />

Feeding specializations First, we assumed that all<br />

birds selected one prey species, with the exception that<br />

they could take a mixture of Scrobicularia <strong>and</strong> Macoma.<br />

This was certainly not the only exception, because<br />

birds also took Cockles <strong>and</strong> Macoma. This was<br />

only observed, however, in summer when Macoma<br />

lived close to the surface at the same <strong>de</strong>pth as Cockles<br />

(Hulscher 1976. Hulsman unpubl.). Consequently, we<br />

were probably wrong in predicting thai birds would<br />

have taken only Macoma in summer 1984 <strong>and</strong> 1985<br />

<strong>and</strong> ignored Cockles.<br />

Another simplification was that all birds should<br />

perform the same prey selection, whereas direct observations<br />

also showed that this was not true. For instance,<br />

one female took only Ragworms among the 29<br />

other colour-b<strong>and</strong>ed birds feeding in the Nes area on<br />

Scrobicularia <strong>and</strong> Macoma (Blomert et al. 1983). Bill<br />

length has a large effect on the prey selection. The last<br />

birds feeding on Scwbicularia in October 1979 <strong>and</strong><br />

the first ones in early spring 1980 were all females<br />

with long bills, because the prey then lived out of reach<br />

of the shorter bill of the males. Also the <strong>de</strong>pletion of<br />

the second year Mya in autumn 1980 was due to predation<br />

by females. The bill length of the birds feeding<br />

on Mya was. on average 1 cm longer than for the birds<br />

feeding at the same time on a cockle bed, just north of<br />

the study area. Hence, it is to be expected that the seasonal<br />

variation in intake rate <strong>and</strong> harvestable <strong>food</strong> supply,<br />

of which the averages has been given in this paper,<br />

were larger for males <strong>and</strong> smaller for females.

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