Stand density and growth of Norway spruce (Picea abies (L.) Karst.) and European beech (Fagus sylvatica L.): Evidence from long-term experimental plots

On the basis of nine Norway spruce (Picea abies (L.) Karst.) and ten European beech (Fagus sylvatica L.) thinning experiments in Germany, for which both residual and removed stock had been registered first during 1870, I scrutinize how moderate and heavy thinning from below (B-, C-grade) affects the production of merchantable volume compared with light thinning (A-grade). In relation to A-grade, cumulative merchantable volume (CV) of B- and C-grade amounts in average to 103-107% in juvenile and to 97-102% in mature Norway spruce stands. The corresponding findings for European beech are 101-106% and 94-102%. CV of individual stands varies between 89% and 130% for Norway spruce and 73% and 155% for European beech (CV of A-grade = 100%). These findings are substantiated by the relation between stand density (SDI) and periodic annual increment (PAI). On the B- and C-grade plots of spruce and beech, respectively, SDI was reduced down to 41-91% and 31-83% of the A-grade. When SDI is reduced in young stands, PAI follows a unimodal curve. Norway spruce's PAI culminates in 109% if SDI is reduced to 59%; European beech's PAI culminates in 123% when density is reduced to 50%. Whereas Norway spruce's growth reacts most positively on thinning under poor site conditions and with increment reduction on favourable sites, European beech behaves oppositely. With stand development the culmination point of the unimodal relation moves towards maximum density, so that in older stands PAI follows the increasing pattern, which is the left portion of a unimodal curve. A model is presented which apparently unifies contradictory patterns of stand density-growth reactions by integrating relative stand density, average tree size and site fertility effects, and makes the findings operable for forest management.