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Past failures of monocultures, caused by wind-throw or insect damages, and ongoing climate change currently strongly stimulate research into mixed-species stands. So far, the focus has mainly been on combinations of species with obvious complementary functional traits. However, for any generalization, a broad overview of the mixing reactions of functionally different tree species in different mixing proportions, patterns and under different site conditions is needed, including assemblages of species with rather similar demands on resources such as light. Here, we studied the growth of Scots pine and oak in mixed versus monospecific stands on 36 triplets located along a productivity gradient across Europe, reaching from Sweden to Spain and from France to Georgia. The set-up represents a wide variation in precipitation (456–1250 mm year−1), mean annual temperature (6.7–11.5 °C) and drought index by de Martonne (21–63 mm °C−1). Stand inventories and increment cores of trees stemming from 40- to 132-year-old, fully stocked stands on 0.04–0.94-ha-sized plots provided insight into how species mixing modifies stand growth and structure compared with neighbouring monospecific stands. On average, the standing stem volume was 436 and 360 m3 ha−1 in the monocultures of Scots pine and oak, respectively, and 418 m3 ha−1 in the mixed stands. The corresponding periodical annual volume increment amounted to 10.5 and 9.1 m3 ha−1 year−1 in the monocultures and 10.5 m3 ha−1 year−1 in the mixed stands. Scots pine showed a 10% larger quadratic mean diameter (p < 0.05), a 7% larger dominant diameter (p < 0.01) and a 9% higher growth of basal area and volume in mixed stands compared with neighbouring monocultures. For Scots pine, the productivity advantages of growing in mixture increased with site index (p < 0.01) and water supply (p < 0.01), while for oak they decreased with site index (p < 0.01). In total, the superior productivity of mixed stands compared to monocultures increased with water supply (p < 0.10). Based on 7843 measured crowns, we found that in mixture both species, but especially oak, had significantly wider crowns (p < 0.001) than in monocultures. On average, we found relatively small effects of species mixing on stand growth and structure. Scots pine benefiting on rich, and oak on poor sites, allows for a mixture that is productive and most likely climate resistant all along a wide ecological gradient. We discuss the potential of this mixture in view of climate change. [ABSTRACT FROM AUTHOR]

• Analysis of drought response patterns in mixed and monospecific stands along a wide ecological gradient across Europe. • Oak exhibits a higher average growth resistance and resilience than Scots pine. • Species mixing increases average drought resistance of Scots pine and resistance and resilience of oak. • Drought responses in mixtures and monocultures are dependent on tree size, site water supply and fertility. Tree species mixing has been widely promoted as a promising silvicultural tool for reducing drought stress. However, so far only a limited number of species combinations have been studied in detail, revealing inconsistent results. In this study, we analysed the effect of mixing Scots pine and oak (pedunculate oak and sessile oak) trees on their drought response along a comprehensive ecological gradient across Europe. The objective was to improve our knowledge of general drought response patterns of two fundamental European tree species in mixed versus monospecific stands. We focused on three null hypotheses: (HI) tree drought response does not differ between Scots pine and oak, (HII) tree drought response of Scots pine and oak is not affected by stand composition (mixture versus monoculture) and (HIII) tree drought response of Scots pine and oak in mixtures and monocultures is not modified by tree size or site conditions. To test the hypotheses, we analysed increment cores of Scots pine and oak, sampled in mixed and monospecific stands, covering a wide range of site conditions. We investigated resistance (the ability to maintain growth levels during drought), recovery (the ability to restore a level of growth after drought) and resilience (the capacity to recover to pre-drought growth levels), involving site-specific drought events that occurred between 1976 and 2015. In monocultures, oak showed a higher resistance and resilience than Scots pine, while recovery was lower. Scots pine in mixed stands exhibited a higher resistance, but also a lower recovery compared with Scots pine in monocultures. Mixing increased the resistance and resilience of oak. Ecological factors such as tree size, site water supply and site fertility were found to have significant effects on the drought response. In the case of Scots pine, resistance was increased by tree size, while recovery was lowered. Resistance of oak increased with site water supply. The observed mixing effect on the tree drought response of Scots pine and oak was in some cases modified by the site conditions studied. Positive mixing effects in terms of resistance and resilience of oak increased with site water supply, while the opposite was found regarding recovery. In contrast, site fertility lessened the positive mixing effect on the resistance of Scots pine. We hypothesise that the observed positive mixing effects under drought mainly result from water- and/or light-related species interactions that improve resource availability and uptake according to temporal and spatial variations in environmental conditions. [ABSTRACT FROM AUTHOR]