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

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Table 3. Three-Way analysis Of variance to lest the effect of the<br />

weight ol ihe total siphon (4-6, ...22-24 mg), ihe experimental conditions<br />

(0. 1-9. 10-15 mg removed), <strong>and</strong> time (Day 0... 5) on the dry<br />

weighl ol the body In = 911: see also Fig. 7.<br />

DEPTH AND SIPHON CROPPING IN SCROBICULARIA<br />

R\ • 1 P<br />

(a) Siphon weighl 26.7 0.001<br />

(b) Exp. condition 6.4 0.026<br />

(c) Time 4.1 0.435<br />

(aix(bi 1 1 (I.S47<br />

(ai xici 4.3 0.977<br />

(b) > 3.2 0.433<br />

Table 3). The animals losing < 9 mg were pooled be­<br />

cause the regression of body weight on siphon weight<br />

was the same. The body weight was relatively low,<br />

however, in the bivalves from which > 10 mg of the<br />

siphon was removed. All animals were alive when they<br />

were dug up 1 to 5 days after the siphon had been removed.<br />

These results agree with the data collected by<br />

Hodgson (1982a) who showed thai the energetic cost<br />

of regeneration was low when 15% ot me siphon was<br />

removed but when 5(W< of more was cut off. regeneration<br />

took place at the expense of the body condition.<br />

Some animals in his experiment died, but no information<br />

is given about survival <strong>and</strong> siphon loss.<br />

n= 2 5 4 6 1 1<br />

'00<br />

30<br />

ۥ 60<br />

survival<br />

20<br />

0<br />

•<br />

i<br />

5<br />

. ,i<br />

10 15<br />

i<br />

20<br />

amount of siphon cut (mg)<br />

Fig. 8. Survival of Scrobicularia (37 mm) kept 2 weeks in an aquarium,<br />

in relation to the amounl of ihe siphon removed I after Straal.<br />

Wanink & Zwarts unpubl. dala).<br />

91<br />

Direct evi<strong>de</strong>nce on the effect of siphon amputation<br />

on subsequent survival is available from an experiment<br />

where Scrobicularia were held in a small aquarium<br />

filled with 20 em mud <strong>and</strong> sea water for 14 days, without<br />

providing <strong>food</strong>. All animals from which 13 to 19<br />

mg of the siphon weight has been removed w ere moribund<br />

or <strong>de</strong>ad, whilst all animals remained in perfect<br />

condition when 0 to 6 mg was taken (Fig. 8).<br />

Siphon weight, burying <strong>de</strong>pth <strong>and</strong> feeding range<br />

The relationship between <strong>de</strong>pth <strong>and</strong> siphon weight<br />

(Fig. 9A. redrawn from Fig. 5). can be used to calculate<br />

the weight per cm of siphon from the upper edge<br />

of the shell to the surface, assuming that Scrobicularia<br />

have to reach the surface (Fig. 9B). It shows that freeliving<br />

animals with a large siphon live at a <strong>de</strong>pth that<br />

allows them 3 mg per cm. Of more interest is the minimum<br />

value: animals with a short siphon, although<br />

they minimize the distance to the surface, only have<br />

available about I mg per cm. Assuming that the maximum<br />

a feeding siphon can be stretched is 1 or 1.5 mg<br />

per cm. it is possible to estimate the potential reach of<br />

the inhalant siphon during surface feeding (Fig. 9C).<br />

Linke (1939). Green (1967. 1968). <strong>and</strong> Hughes<br />

(1969) have given extensive <strong>de</strong>scriptions of the feeding<br />

behaviour of Scrobicularia. but more <strong>de</strong>tailed research<br />

is necessary to confirm that the range on the<br />

surface over which the animals can reach really <strong>de</strong>pends<br />

on siphon weight <strong>and</strong> burying <strong>de</strong>pth, as <strong>de</strong>picted<br />

in Fig. 9C. The mo<strong>de</strong>l presented in Fig. 9 can. however,<br />

be used to <strong>de</strong>rive some predictions.<br />

One prediction is that about half the population will<br />

feed within a radius of 5 to 10 cm around the burrow.<br />

This can be measured directly, but can also be <strong>de</strong>rived<br />

from the length of the starlike feeding tracks which a<br />

Scrobicularia makes during the sucking of the substratum.<br />

The predicted mean value of 5 to 10 cm agrees<br />

well with observations of a normal feeding range of 3<br />

lo 10 cm (Thamdrup 1935), 4 to 10 cm (Linke 1939)<br />

<strong>and</strong> 5 to 8 cm (Hughes 1969). The maximum feeding<br />

range of 20 cm (Linke 1939) falls just within the predicted<br />

range (Fig. 9C I,<br />

It is also to be expected that animals possessing a<br />

heavier siphon <strong>and</strong> thus a larger feeding space, are in a<br />

better condition than animals with a short siphon. This<br />

was in<strong>de</strong>ed the case in the experimental animals (Fig.<br />

7) as well as in nature: the body weight is 2(1 to 30'-'}

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