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

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20 30 40 50<br />

shell length (mm)<br />

PREY PROFITABILITY AND INTAKE RATE<br />

B<br />

10 50 100<br />

prey weight (mg)<br />

o field<br />

A lab; 2cm <strong>de</strong>ep<br />

Fig. 6. H<strong>and</strong>ling lime as a function of A. prev length or B. prey weighl in .V/v.i in captive Ovsiercaichcrs feeding on prey lying al a <strong>de</strong>pth of 2<br />

Ctn I Wanink &. Zwarts 1996) Of wild birds feeding on clams from which lite llesh is eaten in sim (Zwarts & VVaiiink 19S4. Bunskoeke el al.<br />

I9')d & unpubl.). The regressions are calculated without the lour largest cl.mis since llieir h<strong>and</strong>ling times were unlypicallv long. Ihe h<strong>and</strong>ling<br />

limes as a function ol prey weighl are not significantly different Ip = 0.33 in panel Bl. Resulis ol covariance analyses: A. R-' = OXs) tor prev<br />

lengih: B. R-' = 0.869 for prey weight <strong>and</strong> R- = 0.002 lor ihe two groups: Table I gives more <strong>de</strong>tails.<br />

weight of the flesh extracted from Mya of unknown<br />

size, so more field data were available when h<strong>and</strong>ling<br />

time was plotted against ihe prey weight (Fig. 6B). As<br />

Fig. 6B shows, the relationship between h<strong>and</strong>ling time<br />

<strong>and</strong> prey weight was the same as in the laboratory,<br />

where all the llesh from shallow prey was eaten, <strong>and</strong><br />

field studies, where a variable amount of llesh was extracted<br />

from <strong>de</strong>ep-living prey.<br />

In conclusion, the h<strong>and</strong>ling time of all armoured<br />

prey species increases wilh pre) si/e. An additional,<br />

<strong>and</strong> substantial, part of the variation in h<strong>and</strong>ling time<br />

may be explained by four other variables: (I) <strong>de</strong>pth<br />

from which prey are taken: (2i whether prey are hammered<br />

or stabbed: (3) whether prey are opened in situ<br />

or opened after being pulled up from or off the sediment:<br />

<strong>and</strong> (4) portion of the prey eaten.<br />

H<strong>and</strong>ling time of bivalves in relation to prey<br />

condition<br />

Figures 2-6 show the h<strong>and</strong>ling times as a function of<br />

prey length <strong>and</strong> of prey weight. If most of the h<strong>and</strong>ling<br />

time is spent in eating the llesh. we expect that the h<strong>and</strong>ling<br />

time would <strong>de</strong>pend on the amount of llesh ingested<br />

<strong>and</strong> lhat the close relationship between han­<br />

:m<br />

500<br />

dling nine <strong>and</strong> prey size is due only to the high correlation<br />

between prey si/e <strong>and</strong> llesh weight. On the other<br />

h<strong>and</strong>, if h<strong>and</strong>ling time consists mainly of lime speni in<br />

breaking ihc shell, h<strong>and</strong>ling lime would be primarily<br />

<strong>de</strong>termined by size-related strength ol the shell <strong>and</strong> mn<br />

by the amount of llesh ingested. Prey weight <strong>and</strong> prey<br />

size are so highly correlated, that it is hardly surprising<br />

that it is not possible to tell from Figs. 2-6 whether<br />

h<strong>and</strong>ling time <strong>de</strong>pends on prey weight, <strong>and</strong> thus indirectly<br />

on si/e. or on prey si/e. <strong>and</strong> ihus indirectly on<br />

the amount of flesh to be ingested.<br />

Although llesh weight <strong>and</strong> prey size were highly<br />

correlated when both were plotted on a log-log scale.<br />

the weight variation within each si/e class was huge<br />

enough to investigate whether the amounl of llesh. in<strong>de</strong>pen<strong>de</strong>nt<br />

of prey size, affected the h<strong>and</strong>ling time.<br />

Most of the species preyed upon by Oystercatchers<br />

contained in late winter40*3" less llesh than specimens<br />

of similar si/e in early summer (Chambers & Milne<br />

1979. Zwarts 1991, Zwarts & Wanink 1993. Ens et al.<br />

1996b). Prey condition had been <strong>de</strong>fined as percent<br />

<strong>de</strong>v laiion of the average prey weight, such as obtained<br />

by regressing loglweighn againsi log(si/ei. using all<br />

data given in Figs. 2-6. Multiple regression analyses

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