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. Electronic supplementary material

Tree-ring chronologies were examined to investigate the influence of climate on radial growth of Pinus nigra in southeastern Spain. We addressed whether drought differentially affected the ring-widths of dominant and suppressed trees and if our results supported the hypothesis that, in a Mediterranean climate, suppressed conifer trees suffer greater growth reductions than dominant trees. Climate-growth relationships were analyzed using response and correlation functions, whereas the effect of drought on trees growth was approached by superposed epoch analysis in 10 dry years. A cool, wet autumn and spring, and/or mild winter enhanced radial growth. Latewood was the most sensitive ring section in both kinds of trees and it was primarily influenced by current year precipitations. Earlywood was mostly influenced by climatic conditions previous to the growing season. In general, May was the most influential month. Pinus nigra was shown to be very drought sensitive tree in the study area. Tree-rings in suppressed trees showed lower growth reductions caused by drought than those of dominant trees. However, dominant trees recovered normal growth faster. Dominant trees showed a more plastic response, and suppression appeared to reduce the effect of climate on tree radial growth. Some possible causes for these effects are discussed. Our results support the essential role of the balance between light and moisture limitations for plant development during droughts and show that it is not appropriate to generalize about the way in which suppression affects climate-growth relationship in conifers.

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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.

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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. 89 Keywords 91Temporal variability; mixed-species forests; plant-plant interactions; overyielding; 92 asynchrony; niche complementarity; organizational levels; 93 94

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Quantifying the carbon balance in forests is one of the main challenges in forest management. Forest carbon stocks are usually estimated indirectly through biomass equations applied to forest inventories, frequently considering different tree biomass components. The aim of this study is to develop systems of equations for predicting tree biomass components for the main forest softwood species in Spain: Abies alba Mill., A. pinsapo Boiss., Juniperus thurifera L., Pinus canariensis Sweet ex Spreng., P. halepensis Mill., P. nigra Arn., P. pinaster Ait., P. pinea L., P. sylvestris L., P. uncinata Mill. For each species, a system of additive biomass models was fitted using seemingly unrelated regression. Diameter at the breast height and total height were used as independent variables. Diameter appears in all component models, while tree height was included in the stem component model of all species and in some branch component equations. Total height was included in order to improve biomass estimations at different sites. These biomass models were compared to previously available equations in order to test their accuracy and it was found that they yielded better fitting statistics in all cases. Moreover, the models fulfil the additivity property.We also developed root:shoot ratios in order to determine the partitioning into aboveground and belowground biomass. A number of differences were found between species, with a minimum of 0.183 for A. alba and a maximum of 0.385 for P. uncinata. The mean value for the softwood species studied was 0.265.Since the Spanish National Forest Inventory (NFI) records species, tree diameter and height of sample trees, these biomass models and ratios can be used to accurately estimate carbon stocks from NFI data.Key words: aboveground biomass; belowground biomass; carbon sequestration; root:shoot ratio; softwood species. Resumen Nuevos modelos para estimar la capacidad de fijación de carbono de las coníferas españolasConocer el balance de carbono en los bosques es uno de los principales retos dentro de la gestión forestal. Habitualmente, la estimación de carbono en los bosques se realiza de manera indirecta, mediante la aplicación de modelos de diferentes fracciones de biomasa a los datos de inventario forestal. Para ello, en este estudio se han desarrollado sistemas de ecuaciones para estimar la biomasa forestal de las principales coníferas de los bosques de España: Abies alba Mill., A. pinsapo Boiss., Juniperus thurifera L., Pinus canariensis Sweet ex Spreng., P. halepensis Mill., P. nigra Arn., P. pinaster Ait., P. pinea L., P. sylvestris L., P. uncinata Mill. Se ha usado la metodología de mínimos cuadrados generalizados conjuntos, para el cumplimiento de la aditividad entre componentes. Como variables independientes se utilizaron el diámetro normal y la altura total del árbol. El diámetro aparece en todas las ecuaciones como variable significativa, mientras que la altura aparece también en todos los modelos para la biomasa de fuste y en algunos modelos para las ramas. Con l...