Tree species mixing can increase maximum stand density

Mixed-species stands are on the advance in Europe. They fulfil many functions better than monocultures. Recent papers show that mixed stands can have higher yields, but it remains open whether mixed stands simply grow faster along the same self-thinning lines as pure stands or have higher maximum stand densities. We analyzed the effect of species mixing on maximum density based on triplets of pure and mixed stands at approximately maximum density. Most considered mixtures include Norway spruce (Picea abies (L.) H. Karst.). We show that (i) in mixed stands, maximum density is, on average, 16.5% higher than in neighbouring pure stands, and (ii) species mixtures with Norway spruce exceed densities of pure stands by 8.8%, on average. For individual species mixtures, we find a significant density effect of+29.1% for Norway spruce mixed with European larch (Larix decidua Mill.) and +35.9% for Scots pine (Pinus sylvestris L.) in association with European beech (Fagus sylvatica L.). No significant links with stand variables such as age and mean tree size and site fertility were found. The results indicate that species mixing substantially increases stand density, indicating a higher carrying capacity caused by a higher supply and use efficiency of resources. The implications for inventory, silviculture, and forest modelling are discussed.