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

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PREY SIZE SELECTION AND INTAKE RATE<br />

PREY SIZE SELECTION AND INTAKE RATE<br />

Oystercatchers ignore small bivalves. These prey have a lower probability of being encountered, but this<br />

alone is not sufficient to explain why they are not taken. Small bivalves are not profitable given the time<br />

nee<strong>de</strong>d to open <strong>and</strong> eat them, <strong>and</strong> consequently they allow only low rates of return of energy. The size<br />

rejection threshold is not fixed, but varies according to the average intake rate during feeding. Oystercatchers<br />

increase (heir intake rate in two ways: they reject unprofitable prey <strong>and</strong> they adjust <strong>their</strong> searching<br />

behaviour in or<strong>de</strong>r to increase the encounter rate with ihe more profitable prey. The intake rate <strong>de</strong>pends<br />

predominantly on the prey size taken. When large prey are present, the intake rate is high (3 to 4<br />

mg dry flesh s 1 ) <strong>and</strong> small <strong>and</strong> medium-si/cd bivalves are rejected, due to <strong>their</strong> lower profitability. When<br />

only small prey are available, ihe intake rate <strong>de</strong>creases to 1 mg s '.The daily consumption varies between<br />

36 <strong>and</strong> 50 g dry flesh. This means that even when the birds feed at the average year-round rate of 2 mg<br />

s '. 5 h feeding a day is sufficient to meet <strong>their</strong> normal maximum daily energy expenditure. Ninety-two<br />

percent of the intake rates observed in the field excee<strong>de</strong>d the rate at which the gut of Oystercatchers is<br />

able to process <strong>food</strong> (0.66 mg s '). Since processing rate forces the bird to stop feeding when <strong>their</strong><br />

oesophagus is full (12 g). Oystercatchers must stop feeding for digestive pauses when <strong>their</strong> intake rate is<br />

high. Due to this digestive bottleneck, they have to feed 10 hours a day if there is one feeding period, or<br />

8 hours a day if there are two feeding periods, digestive pauses inclu<strong>de</strong>d, to reach the minimum dailv<br />

consumption of 36 g. required at theriiioneutrality.<br />

Introduction<br />

Do predators take all the individual prey belonging to<br />

one species that they encounter, <strong>and</strong> if not. to what <strong>de</strong>gree<br />

are some prey un<strong>de</strong>r-represented in the diet <strong>and</strong><br />

why? These questions are important in every study of<br />

the interaction between predators <strong>and</strong> <strong>their</strong> fcxxl supply.<br />

This chapter first analyses the <strong>de</strong>gree to which the<br />

selection of prey sizes within one prey species by Oystercatchers<br />

can be un<strong>de</strong>rstcxxl as a passive process in<br />

which a r<strong>and</strong>om I v searching bird takes all the prev it<br />

encounters; this will be referred to as 'passive si/e selection".<br />

As we shall see. however, Oystercatchers reject<br />

certain size classes of prey which they actually encounter,<br />

<strong>and</strong> hence there is also "an active size<br />

selection'. The rules that Oystercatchers obey when selecting<br />

size classes actively, <strong>and</strong> whether these can be<br />

<strong>de</strong>rived from optimal foraging mo<strong>de</strong>ls, are (hen discussed.<br />

Finally, ihe chapter discusses how Oystercatchers<br />

maximize <strong>their</strong> intake rate <strong>and</strong> how the rate at which<br />

<strong>food</strong> can be processed sets a limit to the <strong>food</strong> intake.<br />

The analyses are restricted to five bivalve species:<br />

155<br />

the Edible Mussel Mytilus editlis <strong>and</strong> the Edible<br />

Cockle Cerasto<strong>de</strong>rma edule <strong>and</strong> the three clams: Macoma<br />

balihica. Scrobicularia plana <strong>and</strong> Mya armaria.<br />

There are three main methods far obtaining data on<br />

size selection by Oystercatchers eating these prey<br />

species: recovering opened <strong>and</strong> emptied shells, direct<br />

visual observations <strong>and</strong> size-specific <strong>de</strong>pletion. Shell<br />

recovery is particularly convenient <strong>and</strong> is possible because,<br />

having eaten the flesh, the birds leave the<br />

opened shell behind. These are recognizable as prey by<br />

the damage done to the shell <strong>and</strong>/or by iis position in<br />

ihe substraie. Recendy emptied bivalves also contain<br />

some flesh, sometimes along the mantle edge <strong>and</strong> always<br />

where the adductors are atiaehed on the valves.<br />

The collection of these shells is therefore an easy, reliable<br />

method of obtaining a frequency distribution of<br />

all the size classes taken by Oystercatchers in a certain<br />

area, although there may be a sampling bias in the case<br />

of Mussels (Ens 1982. Cayford 1988). It is thus no coinci<strong>de</strong>nce<br />

that there are more papers on prey size seleciion<br />

in Oystercatchers than there are in all the other<br />

wa<strong>de</strong>r species put together.

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