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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 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-PGmix. 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-PGmix can be used to examine the dynamics of mixed-species stands and determine how they may need to be managed. © 2017 Elsevier B.V.

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

Aim of study: Modelling of forest growth and dynamics has focused mainly on pure stands. Mixed-forest management lacks systematic procedures to forecast the impact of silvicultural actions. The main objective of the present work is to review current knowledge and forest model developments that can be applied to mixed forests. Material and methods: Primary research literature was reviewed to determine the state of the art for modelling tree species mixtures, focusing mainly on temperate forests. Main results: The essential principles for predicting stand growth in mixed forests were identified. Forest model applicability in mixtures was analysed. Input data, main model components, output and viewers were presented. Finally, model evaluation procedures and some of the main model platforms were described. Research highlights: Responses to environmental changes and management activities in mixed forests can differ from pure stands. For greater insight into mixed-forest dynamics and ecology, forest scientists and practitioners need new theoretical frameworks, different approaches and innovative solutions for sustainable forest management in the context of environmental and social changes.

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..

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

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.

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.

Main data are basal area increments by triplet, species composition and year, for the study period 1999-2013. Dataset includes data at community level (stand basal area increment), population level (species basal area increment in mixed and monospecific stands), and individual tree level (basal area increments by core, two cores by tree). Moreover data describing the trees used in the analysis is included.

Evidence of tree regeneration failure of some species in the Iberian Peninsula forests warns us about the impact that the global change may exert on the preservation of Mediterranean forests, such as we know them. Predictions agree about an exacerbation of the summer drought there, acknowledged as the main limiting factor for the recruits’ survival. On the other hand, many studies have also proved the relevant role that local heterogeneity has over the spatial distribution of forest species recruitment by providing safe sites. Therefore, to unravel how climate interacts with local factors over juveniles’ performance seems crucial for the design of successful management strategies that allow facing the global warming. Here, we surveyed the natural recruitment of four dominant tree species in seven mountainous regions in the Iberian Peninsula, along entire elevational ranges as surrogates of their climatic ranges. Two of them have alpine and temperate distributions with populations at their rear edge in the Spanish mountains: Fagus sylvatica and Pinus uncinata; and the other two have a genuine Mediterranean distribution: Quercus ilex and Pinus nigra. Our main goal was to analyze for each species the effect of climate, local factors (i.e. light availability, stand structure and ground cover) and the interactions among them to identify the main drivers leading the regeneration process, assessed in terms of presence, abundance and mean annual growth of juveniles. The results showed different environmental factors determining the recruitment patterns of each species. Nevertheless, they highlighted the pervasive role exerted by both climate and fine scale factors, particularly the co-occurring vegetation on recruits’ abundance, and the light availability on their growth. Moreover, we found some interactions among annual mean temperature and local factors, suggesting that climate and local heterogeneity act hierarchically, i.e. the local conditions may mitigate or exacerbat

© 2015 Geobotanisches Institut ETH, Stiftung Ruebel. Climate change is expected to alter the geographic distribution of many plant species worldwide. However, there is still no clear evidence showing a generalised direction and magnitude of these distribution shifts. Here, we have surveyed, in nine mountainous regions in Spain, an array of tree species along entire elevational ranges, as surrogates of their global climatic ranges, to test for elevational shifts towards cooler locations. We analysed the distribution recruitment patterns of five dominant tree species, recording the abundance and measuring the primary growth of juveniles in 306 plots. Three of the species have a temperate-boreal distribution with populations at their southern edge in the Mediterranean mountain ranges: Pinus sylvestris, Pinus uncinata and Fagus sylvatica; and the other two species have a Mediterranean distribution: Quercus ilex and Pinus nigra. Despite the contrasting phylogenies and biogeographies, we identified a similar pattern in recruitment abundance across species, with an asymmetric distribution of juveniles (more recruits in the middle-upper elevation of their range), but higher annual growths at lower elevations. This survival-growth trade-off at the early recruitment stage may potentially counterbalance at population level the negative effect of global warming on recruit survival at the lower edge of species ranges. These findings suggest a demographic stabilisation process at the early recruitment stage of these tree species, and highlight the importance of considering the different demographic stages across the whole climatic range to understand the effects that climate change may exert on species distributions and population dynamics.

Mechanistic water balance models can be used to predict soil moisture dynamics and drought stress in individual forest stands. Predicting current and future levels of plant drought stress is important not only at the local scale, but also at larger, landscape to regional, scales, because these are the management scales at which adaptation and mitigation strategies are implemented. To obtain reliable predictions of soil moisture and plant drought stress over large extents, water balance models need to be complemented with detailed information about the spatial variation of vegetation and soil attributes. We designed, calibrated and validated a water balance model that produces annual estimates of drought intensity and duration for all plant cohorts in a forest stand. Taking Catalonia (NE Spain) as a case study, we coupled this model with plot records from two Spanish forest inventories in which species identity, diameter and height of plant cohorts were available. Leaf area index of each plant cohort was estimated from basal area using species-specific relationships. Vertical root distribution for each species in each forest plot was estimated by determining the distribution that maximized transpiration in the model, given average climatic conditions, soil attributes and stand density. We determined recent trends (period 1980-2010) in drought stress for the main tree species in Catalonia; where forest growth and densification occurs in many areas as a result of rural abandonment and decrease of forest management. Regional increases in drought stress were detected for most tree species, although we found high variation in stress changes among individual forest plots. Moreover, predicted trends in tree drought stress were mainly due to changes in leaf area occurred between the two forest inventories rather than to climatic trends. We conclude that forest structure needs to be explicitly considered in assessments of plant drought stress patterns and trends over large ge

2014/2015

2014/2015

Silver fir (Abies alba) reaches its southwestern distribution limit (rear edge) in the Pyrenees, where it forms highly fragmented populations threatened by drought-induced die-off. Therefore, we need a better knowledge of regeneration patterns and processes (seed production and dispersal, regeneration niche) of such rear-edge stands to assist their long-term conservation. Seed rain patterns were studied in two structurally contrasting sites: an even-aged pure stand (Las Eras) and an uneven-aged mixed silver fir-beech (Fagus sylvatica) stand (Gamueta) over 8 and 12. years, respectively. Seed production experienced a greater synchrony in years of high seed production than when crops were low, suggesting masting behavior in both sites. Seed inverse modeling methods resulted in dispersal curves differing between sites, with estimates of mean dispersal distances of 9.9 and 21.5. m for the pure and mixed stands, respectively. Long-term sampling of seed dispersal and production was complemented with an analysis of silver fir regeneration niche. Seed and seedling patterns were spatially uncoupled, emphasizing the relevance of microsite filtering of regeneration. Understory light environment was the main factor explaining seedling emergence and mortality. Our study provides a mechanistic basis and a methodological approach for understanding the colonization ability of Pyrenean rear-edge silver fir stands and similar tree populations. © 2014 Elsevier B.V.

In some areas of the Mediterranean basin where the understory stratum represents a critical fire hazard, managing the canopy cover to control the understory shrubby vegetation is an ecological alternative to the current mechanical management techniques. In this study, we determine the relationship between the overstory basal area and the cover of the understory shrubby vegetation for different dominant canopy species (Pinaceae and Fagaceae species) along a wide altitudinal gradient in the province of Catalonia (Spain). Analyses were conducted using data from the Spanish National Forest Inventory. At the regional scale, when all stands are analysed together, a strong negative relationship between mean shrub cover and site elevation was found. Among the Pinaceae species, we found fairly good relationships between stand basal area and the maximum development of the shrub stratum for species located at intermediate elevations (Pinus nigra, Pinus sylvestris). However, at the extremes of the elevationclimatic gradient (Pinus halepensis and Pinus uncinata stands), stand basal area explained very little of the shrub cover variation probably because microsite and topographic factors override its effect. Among the Fagaceae species, a negative relationship between basal area and the maximum development of the shrub stratum was found in Quercus humilis and Fagus sylvatica dominated stands but not in Quercus ilex. This can be due to the particular canopy structure and management history of Q. ilex stands. In conclusion, our study revealed a marked effect of the tree layer composition and the environment on the relationship between the development of the understory and overstory tree structure. More fine-grained studies are needed to provide forest managers with more detailed information about the relationship between these two forest strata.