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

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4<br />

Q) Si<br />

•a<br />

a. 16-<br />

20-<br />

A<br />

predator: Calidris alba<br />

prey: Exorolana linguitrons<br />

r}r , Excirolana kincaidn<br />

y<br />

I A<br />

0.0 0 5 1.0<br />

SIPHON SIZE AND DEPTH IN BENTHIC BIVALVES<br />

predator: Haematopus oslralegus<br />

prey: Scrobicularia plana<br />

)0 0.5 1.0<br />

relative prey risk<br />

predator: Numenius arquata<br />

prey: Mya arenaria<br />

Fig. II. Predation risk as a function of burying <strong>de</strong>pth in three different studies: A. S<strong>and</strong>erling Calidris alba preying on beach crustaceans (Myers<br />

el ul. 1180). B. Ovsicrcaicher Haematopus oslralegus feeding on a benthic bivalve (Wanink .V Zwans 1985) <strong>and</strong> C Curlew Numenius ariliiatii<br />

feeding on benthic bivalve (/.warts A: Wanink 19X41. Prev risk is expressed in relation to the maximal predalion risk of a single size class<br />

i set to I); •*• indicates a shallow prey taken bv a Curlew, but not occurring in ihe sampling programme, resulting in an infinitely large prey risk.<br />

overlying water while the siphon is just at the surface<br />

(Brafield & Newell 1961. Hughes 1969. <strong>de</strong> Wil<strong>de</strong><br />

1975. Earll 1975. Hummel 1985a). Macoma lake <strong>de</strong>posit<br />

al a maximum of 4 lo 6 cm from <strong>their</strong> burrow<br />

(Brafield & Newell 1961. Gilbert 1977) <strong>and</strong> Scrobicularia<br />

up to 10 cm (Thamdrup 1935. Hughes 1969) or<br />

even 20 cm (Linke 1939). Both species might be able<br />

to double <strong>their</strong> burying <strong>de</strong>pth by shifting from <strong>de</strong>posit<br />

feeding to filter feeding. Since filter feeding is limited<br />

to the immersion period <strong>and</strong> <strong>de</strong>posit feeding can occur<br />

during immersion as well as during emersion, a vertical<br />

movement within a tidal cycle might be expected.<br />

as suggested by Yonge 11953). No evi<strong>de</strong>nce, however,<br />

was found for this in our laboratory (unpublished data)<br />

when we measured continuously the burying <strong>de</strong>pth of<br />

individual Macoma with the aid of thin nylon threads,<br />

which connected the shells with a registration apparatus.<br />

It is likely dial the remarkable difference between<br />

the summer <strong>and</strong> winter burying <strong>de</strong>pths oi Macoma <strong>and</strong><br />

Scrobicularia (Figs. 4 <strong>and</strong> 5) can be explained by the<br />

absence of <strong>de</strong>posit feeding in winter (Linke 1939,<br />

Hughes 1969. Hummel 1985a). Individuals with a<br />

106<br />

similar siphon weight live 2-3 times as <strong>de</strong>ep in winter<br />

as in summer (Figs. 7 <strong>and</strong> 8). The siphon weights in<br />

Scrobicularia are about the same in both seasons (Fig.<br />

6D). whereas for Macoma the siphon is 35% heavier<br />

during winter than in summer (Fig. 6C). so that most<br />

individuals of both species are able to live at greater<br />

<strong>de</strong>pths in the non-feeding season (Figs. 4 <strong>and</strong> 5). The<br />

relatively low siphon weight of Macoma during the<br />

summer can be attributed lo heavy siphon cropping by<br />

fishes, crabs <strong>and</strong> shrimps (<strong>de</strong> Vlas 1985). It would be<br />

of interest to know to what <strong>de</strong>gree die occurrence of<br />

suspension feeding during the summer in Macoma<br />

(Hummel 1985a) is caused by the reduction of siphon<br />

size.<br />

The suspension fee<strong>de</strong>rs Mya <strong>and</strong> Cerasto<strong>de</strong>rma remain<br />

at nearly the same <strong>de</strong>pth during the whole year:<br />

Cerasto<strong>de</strong>rma burrow to a 20'7< greater <strong>de</strong>pth in winter<br />

than in summer (Fig. 3) <strong>and</strong> in Mya the difference is<br />

only 10 to 15* (Figs. 2 <strong>and</strong> 9).<br />

Burying <strong>de</strong>pth as a protection against predation<br />

The relationship between burying <strong>de</strong>pth <strong>and</strong> size can<br />

be <strong>de</strong>scribed with an S-curve (Figs. 2-5). Burying

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