Resumen
The aim of this review and synthesis is to illustrate the gradual transformation of needles and cones from litter to humus. Individual needles may follow quite different decomposition pathways, which contributes to a diverse humus structure. In the litter layer (Oi), about 40% of the needles were excavated by special mites that produced slowly decomposable excrements. In the fermentation layer (Oe), needles which happened to be in close contact with fine roots decomposed more rapidly. Cones decomposed slower than needles during the first 3?5 years, so the role of cones in carbon (C) storage may be greater than indicated by their fraction of fresh litter. Over a 13 years period, potassium (K), magnesium (Mg) and phosphorus (P) in cones was released, while the total amount of calcium (Ca), manganese (Mn), iron (Fe) and aluminium (Al) increased strongly. Nitrogen concentration increased but the total nitrogen content remained rather constant. After 13 years, the cones had sunk about 6 cm into the soil and lost 60% of their dry weight but were morphologically intact. A cone monitored for 28 years was fully recognizable and had not yet reached the stable Oa layer. The most inert decomposition products in the Oa layer were fragments of needles and cone scales, microarthropod excrements and chitinous insect remains.