Growth and yield of mixed versus pure stands of Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) analysed along a productivity gradient through Europe (original) (raw)
Related papers
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 0C), and the drought index by de Martonne (28–61 mm 0C-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.
European Journal of Forest Research
Past failures of monocultures, caused by wind-throw or insect damages, and ongoing climate change currently strongly stimulate research into mixed-species stands. So far, the focus has mainly been on combinations of species with obvious complementary functional traits. However, for any generalization, a broad overview of the mixing reactions of functionally different tree species in different mixing proportions, patterns and under different site conditions is needed, including assemblages of species with rather similar demands on resources such as light. Here, we studied the growth of Scots pine and oak in mixed versus monospecific stands on 36 triplets located along a productivity gradient across Europe, reaching from Sweden to Spain and from France to Georgia. The setup represents a wide variation in precipitation (456-1250 mm year −1), mean annual temperature (6.7-11.5 °C) and drought index by de Martonne (21-63 mm °C −1). Stand inventories and increment cores of trees stemming from 40-to 132-year-old, fully stocked stands on 0.04-0.94-ha-sized plots provided insight into how species mixing modifies stand growth and structure compared with neighbouring monospecific stands. On average, the standing stem volume was 436 and 360 m 3 ha −1 in the monocultures of Scots pine and oak, respectively, and 418 m 3 ha −1 in the mixed stands. The corresponding periodical annual volume increment amounted to 10.5 and 9.1 m 3 ha −1 year −1 in the monocultures and 10.5 m 3 ha −1 year −1 in the mixed stands. Scots pine showed a 10% larger quadratic mean diameter (p < 0.05), a 7% larger dominant diameter (p < 0.01) and a 9% higher growth of basal area and volume in mixed stands compared with neighbouring monocultures. For Scots pine, the productivity advantages of growing in mixture increased with site index (p < 0.01) and water supply (p < 0.01), while for oak they decreased with site index (p < 0.01). In total, the superior productivity of mixed stands compared to monocultures increased with water supply (p < 0.10). Based on 7843 measured crowns, we found that in mixture both species, but especially oak, had significantly wider crowns (p < 0.001) than in monocultures. On average, we found relatively small effects of species mixing on stand growth and structure. Scots pine benefiting on rich, and oak on poor sites, allows for a mixture that is productive and most likely climate resistant all along a wide ecological gradient. We discuss the potential of this mixture in view of climate change.
Forest Ecology and Management, 2017
The maximum size-density relationship (MSDR) reflects the boundary site occupancy and the selfthinning 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 contribute to the development of more precise silvicultural guidelines and growth models based on the self-thinning line. Moreover, they are of particular importance in the discussion of growth and the effects of mixing on mixed species stands.
Effect of species proportion definition on the evaluation of growth in pure vs. mixed stands
Forest Systems, 2014
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.
Annals of Forest Science, 2010
Existing growth and yield plots of pure and mixed stands of Norway spruce (Picea abies (L.) H. Karst.) and European beech (Fagus sylvatica L.) were aggregated in order to unify the somewhat scattered sources of information currently available, as well as to develop a sound working hypothesis about mixing effects. The database contains information from 23 long-term plots, covering an ecological gradient from nutrient poor and dry to nutrient rich and moist sites throughout Central Europe.
Forest Ecology and Management, 2021
Mixed forests are suggested as a strategic adaptation of forest management to climate change. Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) are tree species of high economic and ecological value for European forestry. Both species coexist naturally in a large part of their distributions but there is a lack of knowledge on the ecological functioning of mixtures of these species and how to manage such stands. This paper analyses these species' intra-and inter-specific competition, including size-symmetric vs. size-asymmetric competition, and explore the effect of weather conditions on tree growth and competition. We studied basal area growth at tree level for Scots pine and Norway spruce in mixed versus pure stands in 22 triplets of fullystocked plots along a broad range of ecological conditions across Europe. Stand inventory and increment cores provided insights into how species mixing modifies tree growth compared with neighbouring pure stands. Five different competition indices, weather variables and their interactions were included and checked in basal area growth models using a linear mixed model approach. Interspecific size-asymmetric competition strongly influenced growth for both tree species, and was modulated by weather conditions. However, species height stratification in mixed stands resulted in a greater tree basal area growth of Scots pine (10.5 cm 2 year − 1) than in pure stands (9.3 cm 2 year − 1), as this species occupies the upper canopy layer. Scots pine growth depended on temperature and drought, whereas Norway spruce growth was influenced only by drought. Interspecific siteasymmetric competition increased in cold winters for Scots pine, and decreased after a drought year for Norway spruce. Although mixtures of these species may reduce tree size for Norway spruce, our results suggest that this could be offset by faster growth in Scots pine. How inter-specific competition and weather conditions alter tree growth may have strong implications for the management of Scots pine-Norway spruce mixtures along the rotation period into the ongoing climate change scenario.
Forest Ecology and Management, 2021
Mixed-species stands have been found to be more productive than would be expected from the performance of their component species in monocultures due to facilitation and complementarity between species, although these interactions depend on the combination of species present. Our study focuses on monospecific and mixedspecies stands of Scots pine and Norway spruce using 20 triplets established in nine countries along a climatic gradient across Europe. Differences in mean tree and stand characteristics, productivity and stand structure were assessed. Basal area increment in mixed stands was 8% higher than expected while volume increment was only 2% greater. Scots pine trees growing in mixed-species stands showed 11% larger quadratic mean diameter, 7% larger dominant diameter, 17% higher basal area and 25% higher stand volume than trees growing in monospecific stands. Norway spruce showed only a non-significant tendency to lower mean values of diameters, heights, basal area, as well standing volume in mixtures than monocultures. Stand structure indices differed between mixed stands and monocultures of Scots pine showing a greater stratification in mixed-species stands. Furthermore, the studied morphological traits showed little variability for trees growing in monospecific stands, except for diameter at breast height, crown length and crown length ratio. For trees growing in mixed stands, all the morphological traits of the trees were identified as different. Some of these morphological traits were
European Journal of Forest Research, 2013
The mixture of beech (Fagus sylvatica L.) and oak (sessile oak, Quercus petraea (MATT.) LIEBL., and pedunculate oak, Q. robur L.) is of considerable importance in Europe and will probably become even more important under climate change. Therefore, the performance of oak and beech in mixture was compared with the species' growth in pure stands. Data from 37 long-term mixing experiments in Poland, Germany and Switzerland were pooled for analysis of mixing effects on stand productivity and possible interrelationships with mixing portions or site conditions. We found that on average, mixed stands of oak and beech exceeded biomass productivity in pure stands by 30 % or 1.7 t ha-1 year-1 , as the growth of both species was benefitted by the mixture. However, that the interaction actually ranged from facilitation and overyielding on poor sites to underyielding on fertile sites triggered by competition. An empirically derived interaction model showed volume and dry mass growth changing in mixed stands from gains of 50 % to losses of 10 % depending on site conditions. It is concluded that the analysed mixture grows in accordance with the stress-gradient hypothesis and that our results suggest a site-specific relationship between species mixture and biomass productivity. As a consequence, an adequate species mix should result in increased productivity under steady state as well as climate change. Keywords Facilitation Á Competitive reduction Á Competition Á Stress-gradient Á Overyielding Á Mixing effect Á Long-term mixing experiments Á Climate change Communicated by R. Matyssek.
Forests, 2018
Although mixing tree species is considered an efficient risk-reduction strategy in the face of climate change, the conditions where mixtures are more productive than monocultures are under ongoing debate. Generalizations have been difficult because of the variety of methods used and due to contradictory findings regarding the effects of the species investigated, mixing proportions, and many site and stand conditions. Using data from 960 plots of the Swiss National Forest Inventory data, we assessed whether Picea abies (L.) Karst–Fagus sylvatica L. mixed stands are more productive than pure stands, and whether the mixing effect depends on site- or stand-characteristics. The species proportions were estimated using species proportion by area, which depends on the maximum stand basal area of an unmanaged stand (BAmax). Four different alternatives were used to estimate BAmax and to investigate the effect of these differing alternatives on the estimated mixture effect. On average, the mi...
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.