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

est of bod\ behind: an estimated 229c of the prey flesh
can be lost in this way (Zwarts & Wanink 1984).
It would be worthwhile in future studies to measure
the amount of flesh remaining in the opened shell, but
there are two problems. Prey opened by Oystercatchers
may be systematically cleaned by Oystercatchers
or other waders, such as Turnstones Arenaria interpres
(Swennen 1990). Secondly, part of the prey may be
Stolen by gull and crow species (Zwarts & Drent 1981,
Ens & Goss-Custard 1984. Swennen 1990). As such
prey cannot be distinguished from prey eaten by Oysleicatchers,
they will cause an overesiimalion of the
amount of flesh left behind if gulls and crows arc less
adept than Oystercatchers at removing the flesh from
the valves.
Estimating time spent feeding The time spent feeding
is estimated in two ways, but both tend to overestimale
the percentage of time actually spent feeding.
When birds are observed for lived intervals of 5 or 10
minutes, birds should be chosen ai random, but observers
will be inclined lo start with a feeding, and not
a resting, bird. Inevitably this overestimates the feeding
activity when such observations are used to estimate
the time speni feeding, lor instance, the feeding
activity of Oystercatchers on Leatherjackets Tipula
paludosa in a grassland from sunrise to sunset was estimated
as 83.5% from birds observed for 15 min periods,
but 59.1% from group scans made every 15 min
(Veenstra 1977). Most studies overcome this difficulty
by determining feeding activity from regular scans of
feeding and non-feeding birds, but this method may
also be biased because birds may leave the feeding
area to rest (Brown & O'Connor 1974. Zwarts et al.
1990).
Estimating total time on the feeding area The estimate
of the time spent on the feeding area by the average
bird is too high when the feeding duration is based
on the time birds are present on the feeding site, because
individual birds may arrive later and/or leave
earlier than the average (Ens & Goss-Custard 1984).
The feeding time of individual birds was 1 h less than
the 4 to 6 h estimated across all birds (Zwarts et al.
1996d). The difference is of course much larger during
the breeding season when individual birds feed in
bouts of only 1 h and spend, in total, only 10 to 30% of
the available low water period actually on the feeding
area (Ens etal. 1996a).
INTAKE RATE AND PROCESSING RATE IN OYSTERCATCHER
218
Can high food intakes be explained as estimation
errors?
To what extent can the extremely high crude intake
rales in some studies plotted in Fig. 2C be attributed to
these five types of error? The studies which estimated
an extremely high consumption will be discussed in
turn.
(1) Heppleston (1971) made all his measurements
between 2.5 h before and after dead low water. He
warned that extrapolation to the entire exposure time
would overestimate the total consumption, because the
mussel bed was exposed for 11 h and the birds hardly
fed over the last lew hours. However, as this was not
quantified, it could not be taken into account.
I2I Goss-Custard (1977) feared that his consumption
estimate was too high because his shell-collections
may have been biased towards large Cockles.
Nor did he take into account that Oystercatchers did
not eat all the flesh from the Cockles.
(3) Zwarts & Wanink (1984) studied Oystercatchers
feeding on small Mya in autumn. The birds used a
mixture of three techniques: the birds either only
grasped ihe siphon, or they ate the whole prey in situ or
they lifted the prey to the surface. Although we know
now (Hulscher ei al. 1996. Wanink & Zwarts 1996)
lhai larger prey are lifted more often than small ones,
that study implicitly assumed that Mya found on the
surface were representative of all si/e classes taken, independent
of the feeding technique used. Since the
majority of prey were eaten in situ, the collection of
prey lifted to the surface, probably caused the average
si/e of Mya to be overestimated. Most clams in the
mud were 16 to 30 mm long. The average length of the
prey on offer was 22.8 mm, compared with 28.2 mm
for the prey collected from the surface, a difference of
5.4 mm. Oystercatchers removed, in total. 80% of the
prey over the months of observation. As Mya do not
grow in autumn and winter, we would expect the size
of the sampled prey to have gradually decreased over
ihe season if ()v stercatchers only took the larger prey.
The extensive sampling programme showed, however,
that the average size of the prey remained exactlv the
same, which implies that the frequency distribution of
the prey taken did not differ from that on offer. Consequently,
the prey weight was overestimated by 30%:
the average prey taken was not 28.2 mm and 65.4 mg.
but 22.8 mm and 50 mg. This error alone reduces ihe