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soft f<strong>or</strong> quite some time. It has 2-5 year classes of older needles. Nutritional content of the new needles is high: total N<br />

reaches 4.5% dwt, carbohydrate levels 250 nanomoles/g dwt (Jensen 1991). When needles get older, nitrogen levels drop to<br />

around 1%, whereas carbohydrate levels increase somewhat (Table 1).<br />

Table 1.--Concentrations of nutrients and secondary compounds in needles and male flowers of spruce, Picea abies.<br />

Total Amino Shikimic Quinic Pinitol Hexoses Inositol Cathecin Total<br />

Plant .part Nitrogen acids acid acid Phenolics<br />

% ................... n moles/mg dwt ...................<br />

Male flowers 5 _900 29 213 134 575 18 4<br />

Old top needles 1.2 _450 224 52 61 390 7 12 68<br />

New top needles 4 1,t20 108 625 133 248 17 3 35<br />

In Europe a substantial number of insect species are flush-feeders on spruce although some of them also consume<br />

older foliage, especially late in year <strong>or</strong> at outbreak densities (Table 2). This situation mainly applies to Lepidoptera although<br />

certain sawfly species also are flush-feeders. Another group feeds exclusively on older needles and is mainly comprised of<br />

Hymenoptera. In general, most spruce flush-feeders (e.g., L. monacha) perf<strong>or</strong>m substantially better when reared on the new,<br />

nutritious needles than on the old ones. Most newly hatched larvae die when put on old needles probably because their<br />

mandibles are not strong enough to cope with the hard needles (Jensen 1992). On the other hand, if third instar larvae are put<br />

on old needles, they survive and perf<strong>or</strong>m reasonably well. Preference indices f<strong>or</strong> new needles (Table 3) decrease with instar<br />

number, indicating that larger larvae do not require young needles.<br />

Table 2.--Needle age preferences of insect larvae feeding on N<strong>or</strong>way spruce, Picea abies.<br />

Needle age class<br />

Insect species Newly flushed Current Old<br />

Lepidoptera<br />

Parasyndemis histrionana * (*) _'<br />

Eana argentata * * (*)<br />

Epinotia tedella (*) *<br />

Epinotia nanana * * (*)<br />

Epinotia pygmaea *<br />

Zeiraphera ratzeburgiana *<br />

Orgya antiqua * * (*)<br />

Lymantria monacha * * (*)<br />

Hymenoptera<br />

Cephalcia abietis *<br />

Cephalcia arvensis * *<br />

Gilpinia hercyniae *<br />

Pristiph<strong>or</strong>a abietina *<br />

Pristiph<strong>or</strong>a ambigua *<br />

Pristiph<strong>or</strong>a saxeseni * (*)<br />

Pachynematus scutellatus * * (*)<br />

" * indicates unequivocal primary preferences, (*) indicates secondary preference, often by late instars.<br />

105

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