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When tree‐species mixtures are more productive than monocultures, higher light absorption is often suggested as a cause. However, few studies have quantified this effect and even fewer have examined which light‐related interactions are most important, such as the effects of species interactions on tree allometric relationships and crown architecture, differences in vertical or horizontal canopy structure, phenology of deciduous species or the mixing effects on tree size and stand density. In this study, measurements of tree sizes and stand structures were combined with a detailed tree‐level light model (Maestra) to examine the contribution of each light‐related interaction on tree‐ and stand‐level light absorption at 21 sites, each of which contained a triplet of plots including a mixture and monocultures ofFagus sylvaticaandPinus sylvestris(63 plots). These sites were distributed across the current distribution of these species within Europe. Averaged across all sites, the light absorption of mixtures was 14% higher than the mean of the monocultures. At the whole community level, this positive effect of mixing on light absorption increased as canopy volume or site productivity increased, but was unrelated to climate. At the species population or individual tree levels, the mixing effect on light absorption resulted from light‐related interactions involving vertical canopy structure, stand density, the presence of a deciduous species (F. sylvatica), as well as the effects of mixing on tree size and allometric relationships between diameter and height, crown diameter and crown length. The mixing effects on light absorption were only correlated with the mixing effects on growth forP. sylvestris, suggesting that the mixing effects on this species were driven by the light‐related interactions, whereas mixing effects onF. sylvaticaor whole community growth were probably driven by non‐light‐related interactions. Synthesis. The overall positive effect of mixing on

1. When tree‐species mixtures are more productive than monocultures, higher light absorption is often suggested as a cause. However, few studies have quantified this effect and even fewer have examined which light‐related interactions are most important, such as the effects of species interactions on tree allometric relationships and crown architecture, differences in vertical or horizontal canopy structure, phenology of deciduous species or the mixing effects on tree size and stand density. 2. In this study, measurements of tree sizes and stand structures were combined with a detailed tree‐level light model (Maestra) to examine the contribution of each light‐related interaction on tree‐ and stand‐level light absorption at 21 sites, each of which contained a triplet of plots including a mixture and monocultures of Fagus sylvatica and Pinus sylvestris (63 plots). These sites were distributed across the current distribution of these species within Europe. 3. Averaged across all sites, the light absorption of mixtures was 14% higher than the mean of the monocultures. At the whole community level, this positive effect of mixing on light absorption increased as canopy volume or site productivity increased, but was unrelated to climate. At the species population or individual tree levels, the mixing effect on light absorption resulted from light‐related interactions involving vertical canopy structure, stand density, the presence of a deciduous species (F. sylvatica), as well as the effects of mixing on tree size and allometric relationships between diameter and height, crown diameter and crown length. 4. The mixing effects on light absorption were only correlated with the mixing effects on growth for P. sylvestris, suggesting that the mixing effects on this species were driven by the light-related interactions, whereas mixing effects on F. sylvatica or whole community growth were probably driven by non-light-related interactions. 5. Synthesis. The overall positive effect

Research into mixed-forests has increased substantially in the last decades but the extent to which the new knowledge generated meets practitioners’ concerns and is adequately transmitted to them is unknown. Here we provide the current state of knowledge and future research directions with regards to 10 questions about mixed-forest functioning and management identified and selected by a range of European forest managers during an extensive participatory process. The set of 10 questions were the highest ranked questions from an online prioritization exercise involving 168 managers from 22 different European countries. In general, the topics of major concern for forest managers coincided with the ones that are at the heart of most research projects. They covered important issues related to the management of mixed forests and the role of mixtures for the stability of forests faced with environmental changes and the provision of ecosystem services to society. Our analysis showed that the current scientific knowledge about these questions was rather variable and particularly low for those related to the management of mixed forests over time and the associated costs. We also found that whereas most research projects have sought to evaluate whether mixed forests are more stable or provide more goods and services than monocultures, there is still little information on the underlying mechanisms and trade-offs behind these effects. Similarly, we identified a lack of knowledge on the spatio-temporal scales at which the effects of mixtures on the resistance and adaptability to environmental changes are operating. Our analysis may help researchers to identify what knowledge needs to be better transferred and to better design future research initiatives meeting practitioner’s concerns.

An increasing amount of research is focusing on comparing productivity in monospecific versus mixed stands, although it is difficult to reach a general consensus as mixing effects differ both in sign (over-yielding or under-yielding) and magnitude depending on species composition as well as on site and stand conditions. While long-term experimental plots provide the best option for disentangling the mixing effects, these datasets are not available for all the existing mixtures nor do they cover large gradients of site factors. The objective of this study was to evaluate the effects and uncertainties of tree species mixing on the productivity of Scots pine–European beech stands along the gradient of site conditions in Europe, using models developed from National and Regional Forest Inventory data. We found a positive effect of pine on beech basal area growth, which was slightly greater for the more humid sites. In contrast, beech negatively affected pine basal area growth, although the effects switched to positive in the more humid sites. However, the uncertainty analysis revealed that the effect on pine at mid- and more humid sites was not-significant. Our results agree with studies developed from a European transect of temporal triplets in the same pine–beech mixtures, confirming the suitability of these datasets and methodology for evaluating mixing effects at large scale.

Summary There is increasing evidence that species diversity enhances the temporal stability (TS) of community productivity in different ecosystems, although its effect at the population and tree levels seems to be negative or neutral. Asynchrony in species responses to environmental conditions was found to be one of the main drivers of this stabilizing process. However, the effect of species mixing on the stability of productivity, and the relative importance of the associated mechanisms, remain poorly understood in forest communities. We investigated the way mixing species influenced the TS of productivity in Pinus sylvestris L. and Fagus sylvatica L. forests, and attempted to determine the main drivers among overyielding, asynchrony between species annual growth responses to environmental conditions, and temporal shifts in species interactions. We used a network of 93 experimental plots distributed across Europe to compare the TS of basal area growth over a 15‐year period (1999–2013) in mixed and monospecific forest stands at different organizational levels, namely the community, population and individual tree levels. Mixed stands showed a higher TS of basal area growth than monospecific stands at the community level, but not at the population or individual tree levels. The TS at the community level was related to asynchrony between species growth in mixtures, but not to overyielding nor to asynchrony between species growth in monospecific stands. Temporal shifts in species interactions were also related to asynchrony and to the mixing effect on the TS. Synthesis. Our findings confirm that species mixing can stabilize productivity at the community level, whereas there is a neutral or negative effect on stability at the population and individual tree levels. The contrasting findings regarding the relationships between the temporal stability and asynchrony in species growth in mixed and monospecific stands suggest that the main driver in the stabilizing process may

Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) dominate many of the European forest stands. Also, mixtures of European beech and Scots pine more or less occur over all European countries, but have been scarcely investigated. The area occupied by each species is of high relevance, especially for growth evaluation and comparison of different species in mixed and monospecific stands. Thus, we studied different methods to describe species proportions and their definition as proportion by area. 25 triplets consisting of mixed and monospecific stands were established across Europe ranging from Lithuania to Spain in northern to southern direction and from Bulgaria to Belgium in eastern to western direction. On stand level, the conclusive method for estimating the species proportion as a fraction of the stand area relates the observed density (tree number or basal area) to its potential. This stand-level estimation makes use of the potential from comparable neighboring monospecific stands or from maximum density lines derived from other data, e.g. forest inventories or permanent observations plots. At tree level, the fraction of the stand area occupied by a species can be derived from the proportions of their crown projection area or of their leaf area. The estimates of the potentials obtained from neighboring monospecific stands, especially in older stands, were poorer than those from the maximum density line depending on the Martonne aridity index. Therefore, the stand-level method in combination with the Martonne aridity index for potential densities can be highly recommended. The species’ proportions estimated with this method are best approximated by the proportions of the species’ leaf areas. In forest practice, the most commonly applied method is an ocular estimation of the proportions by crown projection area. Even though the proportions of pine were calculated here by measuring crown projection areas in the field, we found this method to u

Key message This data set provides unique empirical data from triplets of Scots pine (Pinus sylvestrisL.) and European beech (Fagus sylvaticaL.) across Europe. Dendrometric variables are provided for 32 triplets, 96 plots, 7555 treesand 4695 core samples. These data contribute to our understanding of mixed stand dynamics.Dataset access athttp//dx.doi.org/10.5061/dryad.8v04m. Associated metadata available athttps//metadata-afs.nancy.inra.fr/geonetwork/apps/georchestra/?uuid=b3e098ca-e681-4910-9099-0e25d3b4cd52&hl=eng. © 2017, INRA and Springer-Verlag France SAS.

The maximum size-density relationship (MSDR) reflects the boundary site occupancy and the self-thinning line for a given species, being a useful tool in forestry. Studies focusing on the MSDR often do not cover the whole distribution of the studied species, which results in different boundaries for a given species in different regions. A common MSDR is lacking for the increasingly demanded large-scale studies. However, this information is important where silvicultural responses must be prioritized among monospecific stands or where comparisons among maximum and relative stand densities between and within species are required. For the purposes of this study, we used data from 9911 sample plots located in Scots pine and European beech monospecific stands. Both of these species are of considerable importance and widely distributed throughout Europe. The data came from National or Regional Forest Inventories of five European countries (Austria, Germany, France, Spain and Poland) and therefore were distributed across a wide range of climatic conditions. The main aim of this study was to determine whether the MSDR of these species depends on environmental variables and to develop a MSDR model for each species that explain this variability along a climate gradient. The resulting models showed that both parameters of species boundary lines were climate-dependent, but that the pattern of variation differed between species. Hence, the higher the humidity, the steeper the MSDR (more negative exponent) and the higher the intercept for beech, while in the case of pine, the higher the humidity, the straighter the MSDR and the lower the intercept. According to these models, the stand density indices, for a reference diameter of 25 cm, varied with the humidity in a different way for each species. Consequently, the ratio between the two species increases with humidity, although it also depends on stand diameter. These results are in accordance with the yield level theory and could c

1. There is increasing evidence that species diversity enhances the temporal stability (TS) of com_x0002_munity productivity in different ecosystems, although its effect at the population and tree levels seems to be negative or neutral. Asynchrony in species responses to environmental conditions was found to be one of the main drivers of this stabilizing process. However, the effect of species mix_x0002_ing on the stability of productivity, and the relative importance of the associated mechanisms, remain poorly understood in forest communities. 2. We investigated the way mixing species influenced the TS of productivity in Pinus sylvestris L. and Fagus sylvatica L. forests, and attempted to determine the main drivers among overyielding, asynchrony between species annual growth responses to environmental conditions, and temporal shifts in species interactions. We used a network of 93 experimental plots distributed across Europe to compare the TS of basal area growth over a 15-year period (1999–2013) in mixed and monospecific forest stands at different organizational levels, namely the community, population and individual tree levels. 3. Mixed stands showed a higher TS of basal area growth than monospecific stands at the commu_x0002_nity level, but not at the population or individual tree levels. The TS at the community level was related to asynchrony between species growth in mixtures, but not to overyielding nor to asynchrony between species growth in monospecific stands. Temporal shifts in species interactions were also related to asynchrony and to the mixing effect on the TS. 4. Synthesis. Our findings confirm that species mixing can stabilize productivity at the community level, whereas there is a neutral or negative effect on stability at the population and individual tree levels. The contrasting findings regarding the relationships between the temporal stability and asyn_x0002_chrony in species growth in mixed and monospecific stands suggest that the main driver

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

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. © 2016 Elsevier B.V..

The growth and yield of mixed-species stands has become an important topic of research since there are certain advantages of this type of forest as regards functions and services. However, the concepts and methods used to characterize mixed stands need to be understood, as well as harmonized and standardized. In this review we have compiled a set of measures, indices, and methods at stand level to characterize the structure, dynamics, and productivity of mixed stands, and we discuss the pros and cons of their application in growth and yield studies. Parameters for the characterization of mixed stand structure such as stand density, species composition, horizontal (intermingling) and vertical tree distribution pattern, tree size distribution, and age composition are described, detailing the potential as well as the constraints of these parameters for understanding resource capture, use, and efficiency in mixed stands. Furthermore, a set of stand-level parameters was evaluated to characterize the dynamics of mixed stands, e.g. height growth and space partitioning, self- and alien-thinning, and growth partitioning among trees. The deviations and changes in the behaviour of the analysed parameters in comparison with pure stand conditions due to inter-specific interactions are of particular interest. As regards stand productivity, we reviewed site productivity indices, the growth–density relationship in mixed stands as well as methods to compare productivity in mixed versus monospecific stands. Finally, we discuss the main problems associated with the methodology such as up-scaling from tree to stand level as well as the relevance of standardized measures and methods for improving forest growth and yield research in mixed stands. The main challenges are also outlined, especially the need for qualitatively sound data. © 2015, Springer-Verlag Berlin Heidelberg.

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 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 i

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 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 i

Aim of study The aim of this paper is to compare differences in growth per hectare of species in pure and mixed stands as they result from different definitions of species proportions. Area of study We used the data of the Spanish National Forest Inventory for Scots pine and beech mixtures in the province of Navarra and for Scots pine and Pyrenean oak mixtures in the Central mountain range and the North Iberic mountain range. Material and methods Growth models were parameterized with the species growth related to its proportion as dependent variable, and dominant height, quadratic mean diameter, density, and species proportion as independent variables. As proportions we use once proportions by basal area or by stand density index and once these proportions considering the species specific maximum densities. Main results In the pine-beech mixtures, where the maximum densities do not differ very much between species, the mixing effects are very similar, independent of species proportion definitions. In the pine – oak mixture, where the maximum densities in terms of basal area are very different, the equations using the proportions calculated without reference to the maximum densities, result in a distinct overestimation of the mixing effects on growth. Research highlights When comparing growth per hectare of a species in a mixed stand with that of a pure stand, the species proportion must be described as a proportion by area considering the maximum density for the given species, wrong mixing effects could be introduced by inappropriate species proportion definitions. © 2014 Ministerio de Agricultura Pesca y Alimentacion. All rights reserved.

Aim of study We aim at (i) developing a reference definition of mixed forests in order to harmonize comparative research in mixed forests and (ii) briefly review the research perspectives in mixed forests. Area of study The definition is developed in Europe but can be tested worldwide. Material and methods Review of existent definitions of mixed forests based and literature review encompassing dynamics, management and economic valuation of mixed forests. Main results A mixed forest is defined as a forest unit, excluding linear formations, where at least two tree species coexist at any developmental stage, sharing common resources (light, water, and/or soil nutrients). The presence of each of the component species is normally quantified as a proportion of the number of stems or of basal area, although volume, biomass or canopy cover as well as proportions by occupied stand area may be used for specific objectives. A variety of structures and patterns of mixtures can occur, and the interactions between the component species and their relative proportions may change over time. The research perspectives identified are (i) species interactions and responses to hazards, (ii) the concept of maximum density in mixed forests, (iii) conversion of monocultures to mixed-species forest and (iv) economic valuation of ecosystem services provided by mixed forests. Research highlights The definition is considered a high-level one which encompasses previous attempts to define mixed forests. Current fields of research indicate that gradient studies, experimental design approaches, and model simulations are key topics providing new research opportunities. © 2014 Ministerio de Agricultura Pesca y Alimentacion. All rights reserved.

Aim of study: We aim at (i) developing a reference definition of mixed forests in order to harmonize comparative research in mixed forests and (ii) review the research perspectives in mixed forests. Area of study: The definition is developed in Europe but can be tested worldwide. Material and Methods: Review of existent definitions of mixed forests based and literature review encompassing dynamics, management and economic valuation of mixed forests. Main results: A mixed forest is defined as a forest unit, excluding linear formations, where at least two tree species coexist at any developmental stage, sharing common resources (light, water, and/or soil nutrients). The presence of each of the component species is normally quantified as a proportion of the number of stems or of basal area, although volume, biomass or canopy cover as well as proportions by occupied stand area may be used for specific objectives. A variety of structures and patterns of mixtures can occur, and the interactions between the component species and their relative proportions may change over time. The research perspectives identified are (i) species interactions and responses to hazards, (ii) the concept of maximum density in mixed forests, (iii) conversion of monocultures to mixed-species forest and (iv) economic valuation of ecosystem services provided by mixed forests. Research highlights: The definition is considered a high-level one which encompasses previous attempts to define mixed forests. Current fields of research indicate that gradient studies, experimental design approaches, and model simulations are key topics providing new research opportunities., Aim of study: We aim at (i) developing a reference definition of mixed forests in order to harmonize comparative research in mixed forests and (ii) review the research perspectives in mixed forests. Area of study: The definition is developed in Europe but can be tested worldwide. Material and Methods: Review of existent definitions of mixed forests based and literature review encompassing dynamics, management and economic valuation of mixed forests. Main results: A mixed forest is defined as a forest unit, excluding linear formations, where at least two tree species coexist at any developmental stage, sharing common resources (light, water, and/or soil nutrients). The presence of each of the component species is normally quantified as a proportion of the number of stems or of basal area, although volume, biomass or canopy cover as well as proportions by occupied stand area may be used for specific objectives. A variety of structures and patterns of mixtures can occur, and the interactions between the component species and their relative proportions may change over time. The research perspectives identified are (i) species interactions and responses to hazards, (ii) the concept of maximum density in mixed forests, (iii) conversion of monocultures to mixed-species forest and (iv) economic valuation of ecosystem services provided by mixed forests. Research highlights: The definition is considered a high-level one which encompasses previous attempts to define mixed forests. Current fields of research indicate that gradient studies, experimental design approaches, and model simulations are key topics providing new research opportunities.

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 inte

Although they were originally introduced for different purposes, forest models are often used today for scenario development, which includes forest production as one aspect of forest development. However, studies using an independent data set to compare different simulators are rarely found. In this study a subset of National Forest Inventory data for the whole of Austria was compared to simulations of the biogeochemistry model Biome-BGC, the hybrid gap model PICUS and a climate sensitive version of the growth and yield model PrognAus. The models were used to simulate the development of approximately 70 forest inventory sample plots over a period of 15 years. The study focussed on the models' sensitivity to varying environmental conditions; thus, the comparison was based on the mean current annual volume increment per hectare. All models showed a significant average deviation from the inventory (over- or under-estimation). The estimated year-to-year variation was best reproduced by PICUS. However, the 15 year growth trend was also shown by Biome-BGC and PrognAus. Potential model users interested in relating mean current annual volume increment to climate data will need to weigh accuracy against applicability when choosing among these models.

In this study, we present estimated ranges in carbon (C) sequestration per kg nitrogen (N) addition in above-ground biomass and in soil organic matter for forests and heathlands, based on: (i) empirical relations between spatial patterns of carbon uptake and influencing environmental factors including nitrogen deposition (forests only), (ii) 15N field experiments, (iii) long-term low-dose N fertilizer experiments and (iv) results from ecosystem models. The results of the various studies are in close agreement and show that above-ground accumulation of carbon in forests is generally within the range 15–40 kg C/kg N. For heathlands, a range of 5–15 kg C/kg N has been observed based on low-dose N fertilizer experiments. The uncertainty in C sequestration per kg N addition in soils is larger than for above-ground biomass and varies on average between 5 and 35 kg C/kg N for both forests and heathlands. All together these data indicate a total carbon sequestration range of 5–75 kg C/kg N deposition for forest and heathlands, with a most common range of 20–40 kg C/kg N. Results cannot be extrapolated to systems with very high N inputs, nor to other ecosystems, such as peatlands, where the impact of N is much more variable, and may range from C sequestration to C losses.

In Mediterranean forestry, it is important to improve knowledge about mixed stands dynamics, including their productivity. Previous studies have focused on the interactions between different species (competition, reduction of competition and facilitation) depending on site, species composition and structure. • At the centre of this research are the possible differences between pure and mixed stands of Pinus sylvestris and Quercus pyrenaica in terms of density-growth relationships and volume growth per species. bull; Using data from the second and third Spanish National Forest Inventory (606 plots), volume increment models for these species were fitted. Both species displayed a similar density-growth pattern for pure and mixed stands, with a maximum volume growth at maximum density. Volume increment per occupied area was also found to be greater in mixed stands as opposed to pure stands, suggesting a species interaction with reduced levels of competition in the former. However, the total volume growth was generally lower in mixed stands since the growth rate of oak is much lower. • The results highlight the expedience of favouring P. sylvestris-Q. pyrenaica mixed stands with higher proportions of pine trees in order to gain the benefits of a more complex forest whilst retaining an acceptable level of wood production. © 2009 INRA EDP Sciences.

A logistic model for determining the probability of survival in the next 5-year period of two of the main species in the Mediterranean area in Spain, Mediterranean maritime pine (Pinus pinaster Ait.) and Scots pine (Pinus sylvestris L.), is presented. Data were supplied by the Forest Research Centre of the National Institute of Agricultural and Food Research (INIA) of Spain, consisting of a Permanent Sample Plot network (PSP) covering different stand conditions. In this study, 84 plots for Mediterranean maritime pine and 74 plots for Scots pine were used. Several models were tested. Variables concerning tree size, competition and social position of the tree were introduced in stepwise selection. The ratio diameter at breast height versus quadratic mean diameter and the interaction basal area-site index resulted as the most significant variables in the case of Mediterranean maritime pine. For Scots pine, the most significant variables were site index, DBH-1 and the interaction between basal area of larger trees and the coefficient of variation of the DBH distribution as a surrogate of structure. Results indicate that competition, tree size and social position within the stand are present in the survival process of both species. One-sided competition in highly structured stands for Scots pine and two-sided competition for Mediterranean maritime pine, are shown to be also important. Model evaluation has been done with data from the PSP not used in the fitting process. The area under the Receiver Operating Characteristic curve is 0.86 for Mediterranean maritime pine and 0.85 for Scots pine. The overall predicted rates of survival and mortality are shown for different cut-off points, indicating that the use of random uniform numbers yields better results. Afterwards, the performance of the model in thinned stands is tested for both species. Results for Mediterranean maritime pine show that it is appropriate to use the same model both for unthinned and thinned stands. How