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

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Mytllus edults 4m% -<br />

Nephtys hornbergii — ^ -<br />

FOOD SUPPLY HARVESTABLE BY WADERS<br />

•w<br />

Nereis diversicolor H < •w ro<br />

Cerasto<strong>de</strong>rma edule i- i # S<br />

N<br />

C ^ 8<br />

Macoma balthlca y I • 8<br />

IIS<br />

Scrobicularia plana * •<br />

Mya arenana<br />

-m<br />

s mm Crangon crangon s \<br />

mm Crangon crangon s \<br />

mm s \<br />

mm \<br />

Carcinus maenas<br />

+<br />

21<br />

'<br />

22<br />

' , *<br />

23<br />

*<br />

energy content (kJ-g 1 AFDW)<br />

Fig. 2. Average energ) <strong>de</strong>nsity (kJ g ' AFDW ± SE) Ol ten niver-<br />

tebrates; number of measurements are unhealed. According to a<br />

iine-vvav analysis of variance. Ihe -peek's differ significantly:<br />

Ra=0.05,p=0.027.n=423).<br />

al leasi 20'. to the body weight (e.g. <strong>de</strong> Wil<strong>de</strong> &<br />

Berghuis 1978. Zwarts 1991). so a difference between<br />

the energy value of gametes <strong>and</strong> other flesh would<br />

affect the energy <strong>de</strong>nsity of the entire animal.<br />

However, <strong>de</strong> Wil<strong>de</strong> & Berghuis (1978) found that<br />

gamete production in Macoma would raise the energy<br />

<strong>de</strong>nsity of females <strong>and</strong> lower it for males, since the<br />

energy <strong>de</strong>nsity of eggs was 24.7 kJ g ' <strong>and</strong> of sperm<br />

IS.9 kJ g '. 3 kJ above <strong>and</strong> below the average energy<br />

value of Macoma llesh. respectively. At the population<br />

level, it is thus unlikely that gametogenesis would<br />

cause the energy value of an average Macoma to vary<br />

50 : 50. On the other h<strong>and</strong>, the study of <strong>de</strong> Wil<strong>de</strong> &<br />

Berghuis (I97SI shows how <strong>food</strong> value may vary<br />

between individual prey <strong>and</strong> that predators may be<br />

able to increase <strong>their</strong> rate of energy intake by selecting<br />

female pre\ (see e.g. S/aniawska 1984 for Common<br />

Shrimp Crangon crangon: Zwarts & Blomert 1990 for<br />

Fiddler Crab ilea tangeri).<br />

There were also significant differences in the<br />

energy <strong>de</strong>nsities of ten tidal invertebrates consi<strong>de</strong>red<br />

9<br />

us<br />

f<br />

50<br />

(Fig. 2). Although the worm species had. on average, a<br />

higher energy <strong>de</strong>nsity than ihe bivalves, the highest<br />

energy <strong>de</strong>nsity was found in the Common Mussel<br />

Mytilus edulis. Chambers & Milne (1979) found thai,<br />

in ihe Ythan estuary. E. Scotl<strong>and</strong>, the average energy<br />

<strong>de</strong>nsity differed between Mytilus (22.2 kJ g ' AFDW).<br />

Nereis (21.8 U g '». the Edible Cockle Cerasto<strong>de</strong>rma<br />

edule (20.6 kJ g-i) <strong>and</strong> Macoma (20.0 kJ g 1 ). The<br />

species ranked in exactly the same or<strong>de</strong>r as in fiy 2,<br />

bul the values were in all eases below those found in<br />

our study area.<br />

ln most other studies, estimates of energ) <strong>de</strong>nsit)<br />

are similar to those we found. Using conversion factors<br />

for fat. glycogen <strong>and</strong> protein. Dare & Edwards (1975)<br />

arrived al an average energy <strong>de</strong>nsity for Mytilus oi 23.3<br />

kJ g '. very close to the value given in Fig. 2. The<br />

energ) <strong>de</strong>nsit) measured by Bayne & Worrall (1980)<br />

was slightly higher (24 kJ g '. assuming that ashcontent<br />

was 10%), but Hepplesion (1971) found a<br />

slightly lower value: 22.6 kJ g '. The average value we<br />

found for Macoma is halfway between those given by<br />

Chambers & Milne (1979) (20.0 kJ g ') <strong>and</strong> Beukema<br />

& <strong>de</strong> Bruin (1977) (22.9 kJ g •). The published values<br />

for other bivalves are also similar to ours. Thus.<br />

Swennen (1976) found 21.7 kJ g ' for Cerasto<strong>de</strong>rma,<br />

Hughes (1970b) 21.4 kJ g 1 for Scrobicularia, <strong>and</strong><br />

Edwards & Huebner (1977) <strong>and</strong> Winther & Graj<br />

(1985) found 20.8 <strong>and</strong> 21.7 kJ g*', respectively, for<br />

Mya.<br />

Although a correction of 0.3 kJ g' 1 was ma<strong>de</strong> for<br />

the endothennic reaction of CaCO, (see Methods i. the<br />

energ) <strong>de</strong>nsity oi Carcinus was low. This might be due<br />

to the low energy <strong>de</strong>nsity of the organic component of<br />

the skeleton (Zwarts & Blomert 1990). This explanation<br />

is strengthened by Klein Breteler (1975)<br />

who found an energy <strong>de</strong>nsity of 23 kJ g ' in moulting<br />

Can inns with little skeletal material. The presence of<br />

the skeleton probahlv also <strong>de</strong>presses the energy<br />

<strong>de</strong>nsity of the amphipod Corophium voluiaior where,<br />

according to Chambers & Milne (1979) <strong>and</strong> Boates &<br />

Smith (1979). respectively, the energy <strong>de</strong>nsity is only<br />

19.9 or 20.2 Ug-'.<br />

As fat has a higher energy <strong>de</strong>nsity than proteins <strong>and</strong><br />

carbohydrates. Species differences in biochemical<br />

ci imposition, as well as the amount of skeleton present,<br />

would be expected to cause species differences in<br />

energy <strong>de</strong>nsity. The energy <strong>de</strong>nsitv of Coropfuum is

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