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The productivity and functioning of mixed-species forests often differs from that of monocultures. However, the magnitude and direction of these differences are difficult to predict because species interactions can be modified by many potentially interacting climatic and edaphic conditions, stand structure and previous management. Process-based forest growth models could potentially be used to disentangle the effects of these factors and thereby improve our understanding of mixed forest functioning while facilitating their design and silvicultural management. However, to date, the predicted mixing effects of forest growth models have not been compared with measured mixing effects. In this study, 26 sites across Europe, each containing a mixture and monocultures of Fagus sylvatica and Pinus sylvestris , were used to calculate mixing effects on growth and yield and compare them with the mixing effects predicted by the forest growth model 3-PG mix . The climate and edaphic conditions, stand structures and ages varied greatly between sites. The model performed well when predicting the stem mass and total mass (and mixing effects on these components), with model efficiency that was usually >0.7. The model efficiency was lower for growth or smaller components such as foliage mass and root mass. The model was also used to predict how mixing effects would change along gradients in precipitation, temperature, potential available soil water, age, thinning intensity and soil fertility. The predicted patterns were consistent with measurements of mixing effects from published studies. The 3-PG model is a widely used management tool for monospecific stands and this study shows that 3-PG mix can be used to examine the dynamics of mixed-species stands and determine how they may need to be managed. [ABSTRACT FROM AUTHOR]

The mixing of tree species with complementary ecological traits may modify forest functioning regarding productivity, stability, or resilience against disturbances. This may be achieved by a higher heterogeneity in stand structure which is often addressed but rarely quantified. Here, we use 32 triplets of mature and fully stocked monocultures and mixed stands of Scots pine ( Pinus sylvestris L.) and European beech ( Fagus sylvatica L.) located along a productivity and water availability gradient through Europe to examine how mixing modifies the stand structure in terms of stand density, horizontal tree distribution pattern, vertical stand structure, size distribution pattern, and variation in tree morphology. We further analyze how site conditions modify these aspects of stand structure. For this typical mixture of a light demanding and shade tolerant species we show that (i) mixing significantly increases many aspects of structural heterogeneity compared with monocultures, (ii) mixing effects such as an increase of stand density and diversification of vertical structure and tree morphology are caused by species identity (additive effects) but also by species interactions (multiplicative effects), and (iii) superior heterogeneity of mixed stands over monocultures can increase from dry to moist sites. We discuss the implications for analyzing the productivity, for modelling and for the management of mixed species stands. [ABSTRACT FROM AUTHOR]

• Higher resilience and resistance to drought events in mixed stands compared to monospecifics. • Higher benefits of mixture in conifer-broadleaved stands. • The identity of the species mediates tree response to drought. • Increased resilience to drought in drier sites. • Complexity of the relationship between tree growth and drought confirmed. Despite growing evidence that diverse forests play an important role in ecosystem functioning, ensuring the provision of different ecosystem services, whether such diversity improves their response to drought events remains unclear. In this study, we use a large tree-ring database from thirty case studies across nine European countries and eleven species, covering from Mediterranean to hemiboreal forests, to test if the growth response to site specific drought events that occurred between 1975 and 2015 varied between mixed and monospecific stands. In particular, we quantify how stands resist those specific drought events and recover after them, thus analyzing their resilience. For each drought event and forest stand we calculated resistance, recovery, resilience and relative resilience and we related the variation in these indices between monospecific and mixed stands with type of admixture, tree species identity, site aridity gradient, stand basal area and stand age. We found a large variability among case studies, even for those that share similar species composition and have similar climates. On average, mixed stands showed higher resistance, resilience and relative resilience to drought events than monospecific stands. However, the beneficial effect of mixtures could not be generalized, being greatly modulated by the type of admixture and tree species identity, and depending on site water supply and stand characteristics, such as basal area and age. The increase in resilience in mixtures compared with monocultures was greater on the conifer-broadleaved admixtures, and to a lesser extent in the broadleaved-broadleaved combinations. The observed response patterns to drought largely varied among the eleven studied species, thus revealing the importance of functional traits for understanding a species' response to drought across its distribution range. Along the site aridity gradient, resilience and relative resilience to drought increased in drier sites for both monospecific and mixed stands, with an observed trend towards higher resilience in mixed stands in the drier and hotter sites. Our results confirm the complexity of the relationships found of resistance, recovery, resilience and relative resilience with drought when comparing pure vs mixed stands. [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]