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

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

nasi to the Wad<strong>de</strong>n Sea. mussel beds in the Exe do not<br />

disappear from gales <strong>and</strong> ice. such as occurs in ihc<br />

Wad<strong>de</strong>n Sea iDankers & koelemaij I"**). Obert &<br />

Michaelis 1991). Moreover, since ihe annual recruitment<br />

of Mussels in the Exe is less erratic than in the<br />

Wad<strong>de</strong>n Sea <strong>and</strong> the survival of spat siiongly negatively<br />

<strong>de</strong>nsity-<strong>de</strong>pen<strong>de</strong>nt, ihc <strong>de</strong>nsit) of large Mussels<br />

does not vary much (McGrorty et al. 1990).<br />

Were Oystercalchers <strong>de</strong>pen<strong>de</strong>nt on one or two prey<br />

species in the Wad<strong>de</strong>n Sea. they would not be able to<br />

survive a period longer than one to four years. The<br />

buds iii our study area could only stay alive by switching<br />

regularly from one lo another bivalve species <strong>and</strong><br />

lake, in total, four bivalve species during ihe ten years<br />

of observation (Fig. 14). Nevertheless, ihe intake rales<br />

would be insufficient during four years, <strong>and</strong> ihe birds<br />

had to move to alternative feeding areas. The high water<br />

counts showed thai the total number of Oystercatchers<br />

did nol <strong>de</strong>crease during these years along ihis<br />

part of the Frisian coast <strong>and</strong> the nearby isl<strong>and</strong> Engelsmanplaat<br />

(Zegers & Zwarts unpubl.). Therefore, the<br />

birds must have moved to feeding areas siill within<br />

reach of the same high water roosts. Sightings of<br />

colour-b<strong>and</strong>ed birds showed lhat many of the Nes<br />

birds moved to mussel beds on the lower shore, at 1 to<br />

4 km from the Nes area, ln other words, the birds could<br />

not survive a ten year period if they restricted <strong>their</strong><br />

feeding range to I km : of mudflats. However, by extending<br />

<strong>their</strong> individual feeding ranges in an area of<br />

10-20 km', they could probably find enough fcxxl.<br />

The large majority of Oystercalchers in the Wad<strong>de</strong>n<br />

Sea <strong>de</strong>pend in winter on Cockles <strong>and</strong> Mussels. The<br />

year-to-year variation in ihe ixcurrence of both prey is<br />

very large (Beukema ei al. 1993). Unfortunately for<br />

<strong>their</strong> predators, recruiimenl <strong>and</strong> mortality of both<br />

species is related to the winter temperature, by which<br />

ihe fluctuations of the <strong>food</strong> stock of both species lend<br />

to be synchronized (Beukema et al. 1993). Hence.<br />

()> stercatchers wintering in the Wad<strong>de</strong>n Sea must <strong>de</strong>al<br />

with a varying, <strong>and</strong> sometimes low. fcxxl supply.<br />

Are Oystercatchers limited by <strong>their</strong> <strong>food</strong> supply?<br />

Several prey species contribute to the benthic production,<br />

<strong>and</strong> since each bird species restricts its diet to a<br />

limited number of prey species <strong>and</strong> size classes, each<br />

bird species harvests only a part of ihe lotal production.<br />

Oystercalchers restrict <strong>their</strong> diet mainly to bi­<br />

263<br />

valves. They consumed in the Nes area yearly 12<br />

g in'-'. on average, or 21*Jt of the total yearly elimination<br />

of the live bivalve species combined (Table I).<br />

The predation pressure by Oystercatchers on <strong>their</strong> pi e\<br />

has already often been measured, although usually not<br />

as traction of the annual prey elimination, but as frac<br />

tion of the total biomass being removed over the winter<br />

(see recent reviews by Meire 1993 <strong>and</strong> Goss-Custard<br />

el ul. 1996b). Oy stereatchers are able to remove<br />

between 10 <strong>and</strong> 80% of <strong>their</strong> prey m ., winter, but in<br />

most studies ti is 20 lo 40%. Also this study reveals a<br />

large variation in the predation pressure between prey<br />

species (Tables 1 & 2). <strong>and</strong> between years (Fig. 18). In<br />

S? 0.6 -<br />

i- °<br />

6<br />

E<br />

B 2.5<br />

c<br />

$<br />

2.0k<br />

1.5-<br />

1.0-<br />

10<br />

MILD<br />

20 30 40 50<br />

NORMAL COLD SEVERE<br />

Unsen frost in<strong>de</strong>x<br />

80. ®<br />

^\T2?<br />

- y=exp(l 02 - 0.616x)<br />

0.8-<br />

. r=0.91 p=0.03<br />

j j — i<br />

0.4 0.6 0.8<br />

i<br />

1.0<br />

•<br />

1.2<br />

i—<br />

1.4 1.6<br />

l . _ j _<br />

lowest intake rate (mg-s 1 1.8 2.0<br />

)<br />

Fin. 22. Winter mortality ol Ovstercalchers captured llonrj ihe<br />

i COOSl as a I'unclioii nl A. llie Unsen frost in<strong>de</strong>x <strong>and</strong> B. ihc<br />

predicted lowest intake rale in winter in the Nes area; a selection is<br />

ma<strong>de</strong> for recoveries in January-April. The analysis is based on 3424<br />

colour-b<strong>and</strong>ed Oystercatchers <strong>and</strong> exclu<strong>de</strong>d ihe yearlings: from<br />

/warts,/ ul. (IWbei. The frost in<strong>de</strong>x of Unsen 11991)19 <strong>de</strong>fined as<br />

0.000275V 3 + 0.667y + 1.11 Iz. where \ is the tuiiiibcr ol day l w Ufa<br />

a minimum temperature < 0 "C. <strong>and</strong> y <strong>and</strong> / the number of days with<br />

a maximum temperature < 0 C <strong>and</strong> < -II) '('. resp

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