Mortality reduces overyielding in mixed Scots pine and European beech stands along a precipitation gradient in Europe

Hans Pretzsch, Michael Heym, Torben Hilmers, Andrés Bravo-Oviedo, Shamim Ahmed, Christian Ammer, Admir Avdagić, Kamil Bielak, Felipe Bravo, Gediminas Brazaitis, Marek Fabrika, Vaclav Hurt, Viktor Kurylyak, Magnus Löf, Maciej Pach, Quentin Ponette, Ricardo Ruiz-Peinado, Dejan Stojanovic, Miroslav Svoboda, Barbara WolffTzvetan Zlatanov, Miren del Río

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4 Scopus citations


Many studies show that mixed species stands can have higher gross growth, or so-called overyielding, compared with monocultures. However, much less is known about mortality in mixed stands. Knowledge is lacking, for example, of how much of the gross growth is retained in the standing stock and how much is lost due to mortality. Here, we addressed this knowledge gap of mixed stand dynamics by evaluating 23 middle-aged, unthinned triplets of monospecific and mixed plots of Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) repeatedly surveyed over 6–8 years throughout Europe. For explanation of technical terms in this abstract see Box 1. First, mixed stands produced more gross growth (+10%) but less net growth (−28%) compared with the weighted mean growth of monospecific stands. In monospecific stands, 73% of the gross growth was accumulated in the standing stock, whereas only 48% was accumulated in mixed stands. The gross overyielding of pine (2%) was lower than that of beech (18%). However, the net overyielding of beech was still 10%, whereas low growth and dropout of pine caused a substantial reduction from gross to net growth. Second, the mortality rates, the self- and alien-thinning strength, and the stem volume dropout were higher in mixed stands than monospecific stands. The main reason was the lower survival of pine, whereas beech persisted more similarly in mixed compared with monospecific stands. Third, we found a 10% higher stand density in mixed stands compared with monospecific stands at the first survey. This superiority decreased to 5% in the second survey. Fourth, the mixing proportion of Scots pine decreased from 46% to 44% between the first and second survey. The more than doubling of the segregation index (S) calculated by Pielou index (S increased from 0.2 to 0.5), indicated a strong tendency towards demixing due to pine. Fifth, we showed that with increasing water supply the dropout fraction of the gross growth in the mixture slightly decreased for pine, strongly increased for beech, and also increased for the stand as a whole. We discuss how the reduction of inter-specific competition by thinning may enable a continuous benefit of diversity and overyielding of mixed compared with monospecific stands of Scots pine and European beech.

Original languageEnglish
Article number121008
JournalForest Ecology and Management
StatePublished - 1 Jul 2023


  • Demixing
  • Dropout stem volume
  • Gross and net overyielding
  • Mixed species stands
  • Self- and alien-thinning
  • Stand density
  • Tree mortality


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