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

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INTAKE RATE AND PROCESSING RATE IN OYSTERCATCHER<br />

WHY OYSTERCATCHERS HAEMATOPUS OSTRALEGUS<br />

CANNOT MEET THEIR DAILY ENERGY REQUIREMENTS IN A<br />

SINGLE LOW WATER PERIOD<br />

Captive Oystercatchers consume daily 25-40 g dry flesh or 550-850 kJ. of which thej metabolize 450-<br />

7(K) kJ. Free-living Oystercatchers eat more than captive birds but, contrary to expectation, this is not due<br />

to greater activity costs but to a higher body weight. When body weights are equal, free-living <strong>and</strong> captive<br />

Oystercatchers consume the same amount of <strong>food</strong>. The intake rate of Oystercatchers generally varies<br />

between 1 <strong>and</strong> 3 mg dry flesh s ' feeding, but if non-feeding times are inclu<strong>de</strong>d, the cru<strong>de</strong> intake rate<br />

usually varies between 1 <strong>and</strong> 1.5 mg s '. Extremely high intake rates, above 4 mg s" 1 , are only observed<br />

in birds feeding during a short bout after a long resting period. According to Kersten & Visser (1996a)<br />

such high intake rates cannot be sustained for long, because a maximum of 80 g wet flesh, equivalent to<br />

12 g dry tlesh. can be stored in the digestive tract <strong>and</strong> the processing rate does not exceed 4.4 mg wet<br />

flesh s" 1 or 0.66 mg ash-free dry weight (AFDW) s '. Due to this digestive bottleneck, the birds are forced<br />

to spend much time on the feeding area each day. Since the exposure lime of <strong>their</strong> intertidal feeding areas<br />

is usually 5-6 h, Oystercatchers cannot meet <strong>their</strong> daily energy requirements in a single low water period,<br />

which would often suffice if intake rate was the limiting factor. Fora given length of the feeding period,<br />

the bottleneck mo<strong>de</strong>l predicts the maximum cru<strong>de</strong> intake intake, called CIRm.ix. that can be<br />

achieved, i.e. the Ingliesi intake rate including the non-feeding time. When the birds are able to feed lor<br />

less than 3 h. the achieved cru<strong>de</strong> intake rate usually remains far below this maximum, suggesting dial ihc<br />

rate at which prey are found <strong>and</strong> eaten <strong>de</strong>termines the intake rate. The consumption is also usually less<br />

than would be allowed by digestive constraint when the birds feed for 12 h or longer, because the birds<br />

at thermoneutral conditions do not need more Uian 36 g a day. When the birds spend 3 to 12 h on the<br />

feeding area, the average consumption is usually close to. or below the predicted maximum. However,<br />

in a few cases, the maximum was clearly excee<strong>de</strong>d. These studies do not invalidate the bottleneck mo<strong>de</strong>l,<br />

because there is ample reason to believe that <strong>food</strong> consumption was overestimated. A <strong>de</strong>tailed invesii<br />

gaiion of the many sources of error indicates that <strong>food</strong> consumption is more likely to be overestimated<br />

than un<strong>de</strong>restimated in field studies.<br />

Introduction<br />

The present-day consensus is that birds may fail to collect<br />

a sufficient amount of <strong>food</strong> in the time available, if<br />

they fail to choose correctly the prey species to take or<br />

the place to feed. Recently. Kersten & Visser (1996a)<br />

challenged this view by suggesting that Oystercatchers<br />

cannot process the <strong>food</strong> sufficiently fast during ihe<br />

lime available for digestion, so that the digestion rate is<br />

often a more important constratni on consumption than<br />

the rate al which <strong>food</strong> is ingested. At first sight, this is<br />

a remarkable view as the time available lor digestion<br />

213<br />

will always exceed the time available for feeding.<br />

Thus, internal processing of the <strong>food</strong> may commence<br />

immediately after the first prev item has been ingested<br />

<strong>and</strong> continue long after the incoming ti<strong>de</strong> prevents<br />

feeding. According to Kersten & Visser (1996a). <strong>food</strong><br />

is processed at a constant rate, so thai iniake rales can<br />

only exceed this processing rate during periods when<br />

the digestive tract has not yet been filled to capacity.<br />

The digestive bottleneck hypothesis has important<br />

implications, quite apart from challenging the traditional<br />

emphasis placed on maximization of intake rale<br />

in optimalitv mo<strong>de</strong>ls. It throws, for example, a differ-

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