Mixing of complementary tree species may increase stand productivity, mitigate the effects of drought and other risks, and pave the way to forest production systems which may be more resource-use efficient and stable in the face of climate change. However, systematic empirical studies on mixing effects are still missing for many commercially important and widespread species combinations. Here we studied the growth of Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) in mixed versus pure stands on 32 triplets located along a productivity gradient through Europe, reaching from Sweden to Bulgaria and from Spain to the Ukraine. Stand inventory and taking increment cores on the mainly 60-80 year-old trees and 0.02-1.55 ha sized, fully stocked plots provided insight how species mixing modifies the structure, dynamics and productivity compared with neighbouring pure stands. In mixture standing volume (?12 %), stand density (?20 %), basal area growth (?12 %), and stand volume growth (?8 %) were higher Communicated by Peter Biber. than the weighted mean of the neighbouring pure stands. Scots pine and European beech contributed rather equally to the overyielding and overdensity. In mixed stands mean diameter (?20 %) and height (?6 %) of Scots pine was ahead, while both diameter and height growth of European beech were behind (-8 %). The overyielding and overdensity were independent of the site index, the stand growth and yield, and climatic variables despite the wide variation in precipitation (520-1175 mm year -1 ), mean annual temperature (6-10.5°C), and the drought index by de Martonne (28-61 mm°C -1 ) on the sites. Therefore, this species combination is potentially useful for increasing productivity across a wide range of site and climatic conditions. Given the significant overyielding of stand basal area growth but the absence of any relationship with site index and climatic variables, we hypothesize that the overyielding and overdensity results from several different types of interactions (light-, water-, and nutrient-related) that are all important in different circumstances. We discuss the relevance of the results for ecological theory and for the ongoing silvicultural transition from pure to mixed stands and their adaptation to climate change.
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An estimate of net carbon (C) pool changes and long-term C sequestration in trees and soils was made at more than 100 intensively monitored forest plots (level II plots) and scaled up to Europe based on data for more than 6000 forested plots in a systematic 16 km  16 km grid (level I plots). C pool changes in trees at the level II plots were based on repeated forest growth surveys At the level I plots, an estimate of the mean annual C pool changes was derived from stand age and available site quality characteristics. C sequestration, being equal to the long-term C pool changes accounting for CO 2 emissions because of harvest and forest fires, was assumed 33% of the overall C pool changes by growth. C sequestration in the soil were based on calculated nitrogen (N) retention (N deposition minus net N uptake minus N leaching) rates in soils, multiplied by the C/N ratio of the forest soils, using measured data only (level II plots) or a combination of measurements and model calculations (level I plots). Net C sequestration by forests in Europe (both trees and soil) was estimated at 0.117 Gton yr À1 , with the C sequestration in stem wood being approximately four times as high (0.094 Gton yr À1 ) as the C sequestration in the soil (0.023 Gton yr À1 ). The European average impact of an additional N input on the net C sequestration was estimated at approximately 25 kg C kg À1 N for both tree wood and soil. The contribution of an average additional N deposition on European forests of 2.8 kg ha À1 yr À1 in the period 1960-2000 was estimated at 0.0118 Gton yr À1 , being equal to 10% of the net C sequestration in both trees and soil in that period (0.117 Gton yr À1 ).
Despite the increasing relevance of mixed stands due to their potential benefits; little information is available with regard to the effect of mixtures on yield in forest systems. Hence, it is necessary to study inter-specific relationships, and the resulting yield in mixed stands, which may vary with stand development, site or stand density, etc. In Spain, the province of Navarra is considered one of the biodiversity reservoirs; however, mixed forests occupy only a small area, probably as a consequence of management plans, in which there is an excessive focus on the productivity aspect, favoring the presence of pure stands of the most marketable species. The aim of this paper is to study how growth efficiencies of beech (Fagus sylvatica) and pine (Pinus sylvestris) are modified by the admixture of the other species and to determine whether stand density modifies interspecific relationships and to what extent. Two models were fitted from Spanish National Forest Inventory data, for P. sylvestris and F. sylvatica respectively, which relate the growth efficiency of the species, i.e. the volume increment of the species divided by the species proportion by area, with dominant height, quadratic mean diameter, stocking degree, and the species proportions by area of each species. Growth efficiency of pine increased with the admixture of beech, decreasing this positive effect when stocking degree increased. However, the positive effect of pine admixture on beech growth was greater at higher stocking degrees. Growth efficiency of beech was also dependent on stand dominant height, resulting in a net negative mixing effect when stand dominant heights and stocking degrees were simultaneously low. There is a relatively large range of species proportions and stocking degrees which results in transgressive overyielding: higher volume increments in mixed stands than that of the most productive pure pine stands. We concluded that stocking degree is a key factor in between-species interactions, being the effects of mixing not always greater at higher stand densities, but it depends on species composition.
611. There is increasing evidence that species diversity enhances the temporal stability of 62 community productivity in different ecosystems, although its effect at population and tree 63 levels seems to be negative or neutral. Asynchrony between species was found to be one of 64 the main drivers of this stabilizing process. However, scarce research in this area has been 65 undertaken in forest communities, so determining the effect of species mixing on the stability 66 of forest productivity as well as the identity of the main drivers involved still poses a 67 challenging task. 3. Mixed stands showed a higher temporal stability of basal area growth than monospecific 76 stands at community level, but not at population or individual tree levels. Asynchrony 77 between species growth in mixtures was related to temporal stability, but neither overyielding 78 nor asynchrony between species growth in monospecific stands were linked to temporal 79 stability. Therefore, species interactions modify between-species asynchrony in mixed stands.
80Accordingly, temporal shifts in species interactions were related to asynchrony and to the 81 mixing effect on temporal stability. 4. Synthesis. Our findings confirm that species mixing can stabilize productivity at 83 community level whereas there is a neutral or negative effect on stability at population and 84 individual tree level. The contrasting findings as regards the relationships between temporal 85 stability and species asynchrony in mixed and monospecific stands suggest that the main 86 driver in the stabilizing process is the temporal niche complementarity between species rather 87 than differences in species specific responses to environmental conditions.
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Keywords
91Temporal variability; mixed-species forests; plant-plant interactions; overyielding; 92 asynchrony; niche complementarity; organizational levels; 93 94
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