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Table 3.--Lymantria monacha preference f<strong>or</strong> new needles of spruce, Picea abies.<br />

Instar Preference index n P<br />

I 1.00 + 0.00 20 < 0.001<br />

II 1.00 + 0.00 17 < 0.001<br />

III 0.84 + 0.32 9 < 0.020<br />

IV 0.63 + 0.15 21 < 0.020<br />

V 0.54 + 0.26 21 n.s.<br />

However, as indicated in Table 2, some species actually prefer the old needles, and to such an extent that they will die<br />

if fed the tender, nutritious new needles. An example of <strong>this</strong> is the sawfly, Gilpinia hercyniae, known in N<strong>or</strong>th America as<br />

the European Spruce Sawfly. It is a f<strong>or</strong>mer pest in Canada, but not in Europe. When offered only new foliage, <strong>this</strong> defoliato'r<br />

dies without eating (Jensen 1988). Experiments have shown that one possible deterring compound to G. hercyniae could be<br />

quinic acid, which together with shikimic acid is responsible f<strong>or</strong> much of the acidity of these new needles. Quinic acid is<br />

found in huge amounts in the new needles, but in low concentrations in the old ones. When quinic acid was added to old<br />

(preferred) needles in an experiment (Jensen 1988), G. hercyniae refused to eat after concentrations were increased to the<br />

levels of new needles.<br />

With respect to the phenological window, insects like G. hercyniae, feeding on old needles have no problems. In<br />

contrast, the flush-feeders must find the phenological window. If they hatch very early, even dispersal will not help if the<br />

food resource is not yet present. The only way to survive is to find an alternative food. In the dense monocultures of<br />

European spruce f<strong>or</strong>ests it is not easy to find other host species. However, occasionally, i.e., with 2-7 year intervals flowering<br />

buds might offer <strong>this</strong> alternative. The pollen buds are prime food, being soft, with high nutritional content and rather low<br />

content of secondary compounds (Table 1). Lab<strong>or</strong>at<strong>or</strong>y experiments on newly emerged L. monacha showed that these larvae<br />

are very fond of male flowers. When they hatch early and are reared on branches with only old needles, they die. If male<br />

infl<strong>or</strong>escences are present they survive and grow well (Fig. 1). Once started on flowers, they can eat new and old needles.<br />

Later in the season the larvae may struggle f<strong>or</strong> some time until vegetative buds burst.<br />

0.5 - Weight (g)<br />

0.3 - ""<br />

0.4 - o,,o<br />

0.2 - ,," __"<br />

0.1 - _o",, ,. "<br />

0.02<br />

/<br />

0.04 0.00<br />

_/_/ ,,"<br />

16. May<br />

o Male flowers ->New needles<br />

® Male flowers ->Old needles<br />

Old needles ->Old needles<br />

Figure 1.--Lymantria monacha growth on male flowers and needles of N<strong>or</strong>way spruce (Picea abies).<br />

106

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