waders and their estuarine food supplies - Vlaams Instituut voor de ...
waders and their estuarine food supplies - Vlaams Instituut voor de ...
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 />
total biomass<br />
io.ee ,<br />
harvestable harvestable biomass biomass<br />
+0.68 _<br />
•0.74 _<br />
intake rate<br />
•0.93<br />
bird <strong>de</strong>nsity<br />
. +0.96<br />
+0.95<br />
Fig. 21. Depen<strong>de</strong>nce of bird <strong>de</strong>nsit] on <strong>food</strong> supply <strong>and</strong> predicted<br />
intake rate. The correlations are shown along the arrows: same data<br />
as I ig 20.<br />
Response of Oystercatchers to variation in<br />
harvestable prey biomass<br />
The Nes area was certainly not a marginal feeding<br />
area. The biomass of the <strong>food</strong> supply <strong>and</strong> the feeding<br />
<strong>de</strong>nsity of Oystercatchers were, on average, respectively,<br />
4 <strong>and</strong> 10 times as high as Ihe average for Ihe<br />
tidal flats of the Dutch Wad<strong>de</strong>n Sea as a whole<br />
(Beukema 1976, Zwarts & Wanink 1993). The bird<br />
<strong>de</strong>nsity fluctuated in accordance with the <strong>food</strong> supply,<br />
as shown in Fig. 20A. using the December data from<br />
Fig. lb.<br />
There is a good correlation between bird <strong>de</strong>nsity<br />
<strong>and</strong> "total biomass". r = 0.74. which is actually surprising,<br />
because total biomass is an inaccurate measure of<br />
the fcxxl supply, due to the highly variable fraction that<br />
is unharvestable. Because of this, we expect a better<br />
correlation between bird <strong>de</strong>nsit) <strong>and</strong> the biomass ..|<br />
ihe prey that is both profitable as well as available, <strong>and</strong><br />
this in<strong>de</strong>ed appeared to be the case, r = 0.95 t lig. 20B).<br />
Very likely, this response is caused by the birds acting<br />
on the intake rales that they achieve i lie. HOC). Figure<br />
21 summarizes the causal relationships, as we see<br />
them Two relationships are essential: bird <strong>de</strong>nsity <strong>de</strong>pends<br />
on intake rale, which in turn rate <strong>de</strong>pends on the<br />
harvestable biomass. The high correlation between intake<br />
rale <strong>and</strong> harvestable biomass is not surprising, because<br />
the intake rate has been predicted from the <strong>de</strong>nsity,<br />
si/e <strong>and</strong> burying <strong>de</strong>pth of the prey, three variables<br />
which all contribute to harvestable biomass. It is al<br />
261<br />
ready obvious, however, that many wintering Oystercalchers<br />
only occur in the study area when ol<strong>de</strong>r Cockles<br />
or second year Mya are abundant <strong>and</strong> ihe iniake rale<br />
is > 1.2 mg s*'. In contrast, when ihe wintering birds<br />
have to feed on Scrobicularia or Macoma. hardly any<br />
birds remain to Iced in the .uca because of the extremely<br />
low iniake rale.<br />
The corrcialions show n alongsi<strong>de</strong> the arrow s in Fig.<br />
21, are based upon linear regressions, However, instead<br />
of a linear relationship between bird <strong>de</strong>nsity <strong>and</strong><br />
intake rate (Fig. 20C). we rather expect a J- orS-curve.<br />
No birds should ever feed in the area when the intakeis<br />
< I mg s ; . as we confirmed in this stud) (Fig. 20C).<br />
What will happen above this level will also <strong>de</strong>pend on<br />
what the birds can obtain elsewhere. In other words.<br />
even at a rather high intake rate, few birds may be concentrated<br />
in the area il "the situation elsewhere is even<br />
better. Once intake rales are so high dial the area ranks<br />
among the best in the region, the bird <strong>de</strong>nsity will increase<br />
until all available" birds have been attracted to<br />
the area, after which the <strong>de</strong>nsity will level off al still<br />
higher intake rales because the supplv of recruits dries<br />
up. How man) birds are "available' will <strong>de</strong>pend on the<br />
total number of birds in the region <strong>and</strong> the size ol the<br />
region, which will <strong>de</strong>pend on the size of the home<br />
range of ihe birds. With extreme site fi<strong>de</strong>lity, home<br />
ranges <strong>and</strong> therefore the region will be very small, so<br />
that the pool of 'available' birds is quicklv <strong>de</strong>pleted<br />
<strong>and</strong> very little relationship between bird <strong>de</strong>nsity <strong>and</strong><br />
<strong>food</strong> supply is to be expected when years are com<br />
pared. If. on the other h<strong>and</strong>, home ranges are large <strong>and</strong><br />
variation in loud supplv between years quite extreme.<br />
a high correlation between bird <strong>de</strong>nsit) <strong>and</strong> intake rate<br />
can occur, as was probably the case in our study. The<br />
sightings of the colour-b<strong>and</strong>ed birds certainly proved<br />
thai movements of I to 3 km regularly occurred, although<br />
individuals could stav for inonihs. <strong>and</strong> some<br />
even for years, 00 the same spot of less than 0> ha.<br />
The sile fi<strong>de</strong>lity of the Oystercalchers in our study area<br />
seems intermediate between that in the Exe estuary.<br />
where the mussel beds are extremely stable <strong>and</strong> home<br />
ranges often less than I ha (Goss-C'iistard et al. 1982),<br />
<strong>and</strong> the estuary of the Ribble. where an exceptional])<br />
large spatfall of Cockles caused a massive influx of<br />
Oystercatchers < Suiherl<strong>and</strong> I982d).