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

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June 1981 June 1982<br />

is * N.MUD • Q<br />

10 — r*<br />

e \MUD<br />

o<br />

JC 12 -<br />

a. 1><br />

a<br />

"O<br />

§14 - -<br />

"3<br />

JO<br />

16<br />

\sAND<br />

"<br />

18<br />

" _ • ,t<br />

BURROWING AND FEEDING IN NEREIS<br />

SAND\<br />

,!,,,, - It,<br />

-40 0 +40 +80 -40 0 +40<br />

<strong>de</strong>viation from mean body weight (%)<br />

+80<br />

Fig. 5. .\, mis diversicolor. Burrow <strong>de</strong>pth of worms of 10(7 lo 141<br />

cm living in mud (•) <strong>and</strong> s<strong>and</strong> (O) as function of body condilion in<br />

2 sampling programmes. Mean burrow <strong>de</strong>pth <strong>and</strong> mean body condition<br />

in mud <strong>and</strong> s<strong>and</strong> are indicated by black <strong>and</strong> white arrows, respectively.<br />

A correction has heen ma<strong>de</strong>, .is in Fig 4. to adjust for the<br />

si/c eltect on <strong>de</strong>plh See Table -I lor slatistical analysis.<br />

Figure 4 shows the relationship between clay content<br />

<strong>and</strong> burrow <strong>de</strong>pth. Burrow <strong>de</strong>pth appeared to remain<br />

constant if the clay content increased from 6 to<br />

2(l'». However, in sediments with a clay content of<br />

< 5% (<strong>de</strong>fined above as s<strong>and</strong>) burrow <strong>de</strong>pth increased<br />

when clay content <strong>de</strong>creased from 5 to 2'7c. Nereis are<br />

not found in sediments where the clay content is well<br />

un<strong>de</strong>r 19r (Zwarts 1988b). The relationship between<br />

claj content <strong>and</strong> burrow <strong>de</strong>pth in the s<strong>and</strong>y sediments<br />

\\ as more pronounced in September than in June (Fig.<br />

4). The effect of clay content on burrow <strong>de</strong>pth cannoi<br />

be attributed to other measured variables, such as elevation<br />

(or emersion time 1 .<br />

Burrow <strong>de</strong>pth <strong>and</strong> body condition<br />

The large variation in burrow <strong>de</strong>pth for individuals ul<br />

the same size i Kg, 2) can he explained partly by a difference<br />

in body condition (Fig. 5). The body weight of<br />

<strong>de</strong>ep-living Nereis was on average 2 to 3 limes greater<br />

than thai of individuals of the same si/e class with<br />

shallow burrows. Apparently, worms in good condition<br />

are able to maintain <strong>de</strong>eper burrows.<br />

It is known that body condition varies throughout<br />

Ihe year <strong>and</strong> also differs for WOTmS lnund alone the<br />

tidal gradient (Chambers & Milne 1975b, Esselink et<br />

al. 1989). Figure 5 shows that the bodv condition of<br />

worms living in s<strong>and</strong> was If) to 35% higher than that of<br />

individuals sampled in mud. The effect of clay content<br />

on burrow <strong>de</strong>pth, as <strong>de</strong>scribed in the previous section,<br />

disappears completely in 3 out of 4 sampling programmes<br />

if body condilion is taken into account (Table<br />

4). The <strong>de</strong>eper burrows found in s<strong>and</strong>y sediments<br />

might thus be explained by the average difference in<br />

body weight between individuals inhabiting s<strong>and</strong>y <strong>and</strong><br />

muddy sediments.<br />

There is also a positive relationship between body<br />

condition <strong>and</strong> burrow <strong>de</strong>pth when worms within a<br />

same season are compared, but this effect appears to be<br />

in<strong>de</strong>pen<strong>de</strong>ni of ihe seasonal variation in <strong>de</strong>pth (Fig. 6,<br />

Table 5).<br />

Table 4. Kcsulls of four 2-way .111.ily sc- of variance to test the effect of body coiidiliun <strong>and</strong> clay content on burrow <strong>de</strong>pth in 4 sampling programmes<br />

1 dala lor June <strong>and</strong> July shown in Rg. 5). Since body condilion is related 10 clay contenl. the total variance explained by the 2 main effect*<br />

can be larger than ihc sum of the variances unique to each of the separale laciors<br />

Nov. 1980 April 1981 June I98J Juls 1982<br />

Sonne of 1: ---5S n = 67 n = 9/ n z<br />

s-arialion R>.% P R 2 .% I' /' R>.% /'<br />

Main effects 36.0 0.001 27.2 0.001 54.2 0.001 11.9 0.001<br />

condition 8.0 0.002 25.0 0.001 Mi 0.001 10.4 0.000<br />

cl.iv contenl 7.5 0.020 4.4 0.055 111! 0.X76 0.7 0.118<br />

Two-way interaction n: 0.734 3.7 0.206 0.4 0.718 7 : 0.190<br />

307

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