Mortality Reduces Overyielding in Mixed Scots Pine and European Beech Stands Along a Precipitation Gradient in Europe

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.
The study received funding from the European Union's Horizon 2020 research and innovation programme under grant agreement No952314 and under the Marie Skłodowska-Curie grant agreement No 778322. The first author also wishes to thank the German ScienceFoundation (Deutsche Forschungsgemeinschaft) for funding the project “Structure and dynamics of mixed-species stands of Scots pine and European beech compared with monospecific stands; analysis along an ecological gradient through Europe” (# DFG PR 292/15-1). Felipe Bravo is grateful for Funds by the Junta de Castilla y León through the projects “CLU-2019-01 and CL-EI-2021-05 - iuFOR Institute Unit of Excellence” of the University of Valladolid and the co-financing by the European Regional Development Fund (ERDF “Europe drives our growth”). Miren del Río thanks for the support by the Spanish Ministerio de Ciencia e Innovación (# PID2021-126275OB-C21/C22). Dejan Stojanović thanks the Ministry of Education, Science and Technological Development of the Republic of Serbia for funding. Tzvetan Zlatanov thanks the Ministry of Education and Science of the Republic of Bulgaria (# DO1-405/18.12.2020 LTER-BG).