Mixing effect on volume growth of Fagus sylvatica and Pinus sylvestris is modulated by stand density

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

Mortality reduces overyielding in mixed Scots pine and European beech stands along a precipitation gradient in Europe

Many studies show that mixed species stands can have higher gross growth, or so-called overyielding, compared with monocultures. However, much less is known about mortality in mixed stands. Knowledge is lacking, for example, of how much of the gross growth is retained in the standing stock and how much is lost due to mortality. Here, we addressed this knowledge gap of mixed stand dynamics by evaluating 23 middle-aged, unthinned triplets of monospecific and mixed plots of Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) repeatedly surveyed over 6-8 years throughout Europe. For explanation of technical terms in this abstract see Box 1. First, mixed stands produced more gross growth (+10%) but less net growth (MINUS SIGN 28%) compared with the weighted mean growth of monospecific stands. In monospecific stands, 73% of the gross growth was accumulated in the standing stock, whereas only 48% was accumulated in mixed stands. The gross overyielding of pine (2%) was lower than that of beech (18%). However, the net overyielding of beech was still 10%, whereas low growth and dropout of pine caused a substantial reduction from gross to net growth. Second, the mortality rates, the self- and alien-thinning strength, and the stem volume dropout were higher in mixed stands than monospecific stands. The main reason was the lower survival of pine, whereas beech persisted more similarly in mixed compared with monospecific stands. Third, we found a 10% higher stand density in mixed stands compared with monospecific stands at the first survey. This superiority decreased to 5% in the second survey. Fourth, the mixing proportion of Scots pine decreased from 46% to 44% between the first and second survey. The more than doubling of the segregation index (S) calculated by Pielou index (S increased from 0.2 to 0.5), indicated a strong tendency towards demixing due to pine. Fifth, we showed that with increasing water supply the dropout fraction of the gross growth in the mixture slightly decreased for pine, strongly increased for beech, and also increased for the stand as a whole. We discuss how the reduction of inter-specific competition by thinning may enable a continuous benefit of diversity and overyielding of mixed compared with monospecific stands of Scots pine and European beech.OA-hybri

Implications of reduced stand density on tree growth and drought susceptibility: A study of three species under varying climate

Producción CientíficaA higher frequency of increasingly severe droughts highlights the need for short-term measures to adapt existing forests to climate change. The maintenance of reduced stand densities has been proposed as a promising silvicultural tool for mitigating drought stress. However, the relationship between stand density and tree drought susceptibility remains poorly understood, especially across ecological gradients. Here, we analysed the effect of reduced stand density on tree growth and growth sensitivity, as well as on short-term drought responses (resistance, recovery, and resilience) of Scots pine (Pinus sylvestris L.), sessile oak (Quercus petraea (Matt.) Liebl.), and ponderosa pine (Pinus ponderosa Douglas ex C. Lawson). Tree ring series from 409 trees, growing in stands of varying stand density, were analysed at sites with different water availability. For all species, mean tree growth was significantly higher under low compared with maximum stand density. Mean tree growth sensitivity of Scots pine was significantly higher under low compared with moderate and maximum stand density, while growth sensitivity of ponderosa pine peaked under maximum stand density. Recovery and resilience of Scots pine, as well as recovery of sessile oak and ponderosa pine, decreased with increasing stand density. In contrast, resistance and resilience of ponderosa pine significantly increased with increasing stand density. Higher site water availability was associated with significantly reduced drought response indices of Scots pine and sessile oak in general, except for resistance of oak. In ponderosa pine, higher site water availability significantly lessened recovery. Higher site water availability significantly moderated the positive effect of reduced stand density on drought responses. Stand age had a significantly positive effect on the resistance of Scots pine and a negative effect on recovery of sessile oak. We discuss potential causes for the observed response patterns, derive implications for adaptive forest management, and make recommendations for further research in this field.Eva Mayr-Stihl Stiftung (170124)ERANet SUMFOREST project REFORM- AEI (PCIN-2017-026)German Federal Ministry of Food and Agriculture (2816ERA02S

Modelos de masa para orientar la gestión de Pinus pinea L. en el noreste de España

A yield model was developed to simulate silviculture alternatives for Pinus pinea L. in north-east Spain (Catalonia). The model uses several functions to estimate the main silvicultural parameters at stand level and a disaggregation system to predict diameter distributions. From a network of 75 temporary plots a system of equations to predict stand variables was simultaneously fitted for two stand density types, namely low and high density stands, using the threestage least-squares method (3SLS). The diameter distributions were estimated by the Weibull distribution function using the parameter recovery method (PRM) and the method of moments. Based on this yield model, two silviculture alternatives were simulated for each stand density type and site class, resulting in 16 silviculture scenarios. The yield model and silviculture alternatives offer a management tool and a guide for the sustainable forest management of evenaged Pinus pinea forests in this region. Se ha desarrollado un modelo de masa como base para orientar la gestión de Pinus pinea L. en el noreste de España (Cataluña). El modelo integra diferentes funciones que permiten evaluar las principales variables forestales a nivel de rodal, incluyendo un sistema de desagregación para la estimación de las distribuciones diamétricas. A partir de los datos de una red de 75 parcelas temporales y diferenciando dos tipologías de masa en función de la densidad, masas claras y densas, se ajustaron simultáneamente sistemas de ecuaciones utilizando el método de estimación de mínimos cuadrados en tres etapas. Las funciones obtenidas predicen las principales variables de masa para las dos tipologías. Las distribuciones diamétricas se obtuvieron a partir de la función de distribución de Weibull, utilizando el método de recuperación de parámetros (PRM) y el método de los momentos. Finalmente, se simularon dos alternativas selvícolas para cada tipología de masa (claras, densas) y calidad de estación, resultando 16 escenarios selvícolas. El modelo de masa y las alternativas selvícolas obtenidas se presentan como una herramienta útil para el gestor y una guía para la gestión sostenible de las masas de Pinus pinea en esta región

Variabilidad ambiental de las masas de pino negral y su relación con el índice de sitio

Environmental variability and site productivity relationships, estimated by means of soil-site equations, are considered a milestone in decision making of forest management. The adequacy of silvicultural systems is related to tree response to environmental conditions. The objectives of this paper are to study climatic and edaphic variability in Mediterranean Maritime pine (Pinus pinaster) forests in Spain, and the practical use of such variability in determining forest productivity by means of site index estimation. Principal component analysis was used to describe environmental conditions and patterns. Site index predictive models were fitted using partial least squares and parsimoniously by ordinary least square. Climatic variables along with parent material defined an ecological regionalization from warm and humid to cold and dry sites. Results showed that temperature and precipitation in autumn and winter, along with longitudinal gradient define extreme site qualities. The best qualities are located in warm and humid sites whereas the poorest ones are found in cold and dry regions. Site index values are poorly explained by soil properties. However, clay content in the first mineral horizon improved the soil-site model considerably. Climate is the main driver of productivity of Mediterranean Maritime pine in a broad scale. Site index differences within a homogenous climatic region are associated to soil properties.La relación entre variabilidad ambiental y la productividad de estación, estimada mediante el índice de sitio, es clave en la toma de decisiones en la gestión forestal sostenible, ya que su conocimiento permite adecuar la práctica selvícola a la respuesta de la masa a dicha variabilidad ambiental. Los objetivos de este trabajo son estudiar la variabilidad climática y edáfica de Pinus pinaster en su distribución mediterránea en España y el uso práctico de dicha variabilidad en la determinación de la productividad de la estación mediante la estimación del índice de sitio. Para la descripción de la variabilidad ambiental se realizó un análisis de componentes principales y para la predicción del índice de sitio se optó por una regresión por mínimos cuadrados parciales, y de forma más parsimoniosa, mediante mínimos cuadrados ordinarios. Las variables climáticas, junto al material parental definieron regiones que comprendían estaciones que van de cálidas y húmedas hasta frías y secas. Los resultados mostraron cómo la temperatura media anual, la precipitación en otoño e invierno, junto con un gradiente longitudinal define calidades de estación extremas. Las mejores calidades se encuentran en estaciones cálidas y húmedas mientras que las peores están en estaciones frías y secas. Las variables edáficas explican poca variación del índice de sitio, aunque la inclusión del contenido en arcilla mejora notablemente el modelo. El clima es el precursor de la calidad de estación mientras que diferencias en el índice de sitio en zonas climáticamente homogéneas se asocian a variables edáficas

Changes in structural heterogeneity and stand productivity by mixing Scots pine and Maritime pine

Mixed-species stands have been studied extensively due to their potentially superior productivity, multi-functionality benefits and high ecological value compared to pure stands. The higher structural heterogeneity in mixed stands that can emerge from species interactions could be linked to the relationship between species diversity and ecosystem functions. We tested whether changes in stand structure also occur in mixtures of species with similar traits and whether they explain over-yielding patterns. Based on research with 12 triplets of Scots pine (Pinus sylvestris L.) and Maritime pine (Pinus pinaster Ait.) in the northern Iberian Peninsula (Spain), we provide evidence that species mixing increased structural heterogeneity and may induce over-yielding in mixed-species stands compared to monospecific stands. In this mixture of two light-demanding species, we observed that (i) stand composition influenced the inter-specific crown allometric variation, (ii) structural heterogeneity in mixed stands was caused by both specie-specific traits and species interactions, and (iii) intraspecific and interspecific differences in both crown size plasticity and size-distribution differentiation were associated with the increased relative productivity of mixed stands. We detected that crown complementarity and vertical stratification in the canopy space is a crucial mechanism for enhancing ecosystem productivity in light-demanding species and could be related to light interception and light-use. This work improves our understanding of emerging properties in mixed stands and introduces considerations for properly scaling and tracing mixing effects at individual tree, size distribution and stand levels.Instituto Universitario de Gestión Forestal Sostenibl

Analyzing size-symmetric vs. size-asymmetric and intra-vs. inter-specific competition in beech (Fagus sylvatica L.) mixed stands

In mixed stands, inter-specific competition can be lower than intra-specific competition when niche complementarity and/or facilitation between species prevail. These positive interactions can take place at belowground and/or aboveground levels. Belowground competition tends to be size symmetric while the aboveground competition is usually for light and almost always size-asymmetric. Interactions between forest tree species can be explored analyzing growth at tree level by comparing intra and inter-specific competition. At the same time, possible causes of niche complementarity can be inferred relating intra and inter-specific competition with the mode of competition, i.e. size-symmetric or sizeasymmetric. The aim of this paper is to further our understanding of the interactions between species and to detect possible causes of competition reduction in mixed stands of beech (Fagus sylvatica L.) with other species: pine?beech, oak?beech and fir?beech. To test whether species growth is better explained by size-symmetric and/or size-asymmetric competition, five different competition structures where included in basal area growth models fitted using data from the Spanish National Forest Inventory for the Pyrenees. These models considered either size-symmetry only (Reineke?s stand density index, SDI), size-asymmetry only (SDI of large trees or SDI of small trees), or both combined. In order to assess the influence of the admixture, these indices were introduced in two different ways, one of which was to consider that trees of all species compete in a similar way, and the other was to split the stand density indices into intra- and inter-specific competition components. The results showed that in pine?beech mixtures, there is a slightly negative effect of beech on pine basal area growth while beech benefitted from the admixture of Scots pine; this positive effect being greater as the proportion of pine trees in larger size classes increases. In oak?beech mixtures, beech growth was also positively influenced by the presence of oaks that were larger than the beech trees. The growth of oak, however, decreased when the proportion of beech in SDI increased, although the presence of beech in larger size classes promoted oak growth. Finally, in fir?beech mixtures, neither fir nor beech basal area growth were influenced by the presence of the other species. The results indicate that size-asymmetric is stronger than size-symmetric competition in these mixtures, highlighting the importance of light in competition. Positive species interactions in size-asymmetric competition involved a reduction of asymmetry in tree size-growth relationships

Desfronde y tasa de Descomposición foliar en rebollar-pinar de repoblación con distinto grado de clara

El desfronde representa un factor clave en el funcionamiento de los ecosistemas forestales. Por un lado representa el mayor aporte de nutrientes en sistemas naturales y, por otro, su acumulación en el suelo constituye un banco de elementos esenciales que, por descomposición y mineralización, se integran en el suelo y en los ciclos biogeoquímicos. Este trabajo analiza el efecto de la clara en la producción de desfronde, la eficiencia en el uso del nitrógeno y en la tasa de descomposición de la hojarasca en una masa mixta de Pinus pinaster Ait. procedente de repoblación sobre una masa natural de Quercus pyrenaica Wild. El sitio experimental está situado en los Montes de Toledo y consta de tres tratamientos: control, clara moderada y clara fuerte con tres réplicas en un diseño de cuadrado latino. Los datos de desfronde proceden de recogida mensual en cestos y separación en fracciones para el cálculo de la materia seca. El nitrógeno se analizó en muestras compuestas cada tres meses. Para el estudio de la tasa de descomposición se utilizaron bolsas con mezcla de acículas y hojas, recolectándose y analizándose también cada tres meses. Los resultados muestran picos de desfronde en los meses de verano y en el otoño. La mayor producción de desfronde se observa en las parcelas testigo. La aplicación de las claras no supone grandes diferencias ni en el desfronde ni en la eficencia en el uso del nitrógeno. Por el contrario las claras ralentizan la tasa de descomposición de las hojas de rebollo

Growth response to thinning in a mixed pine-oak stand of Pinus pinaster and Quercus pyrenaica.

During the last decades the interest on mixed stands has increased in forestry due to their potential benefits, especially conifer-broadleaves mixtures. However, more knowledge is need about their dynamic and their response to silvicultural treatments. On the other hand, climate change can affect different species in different ways, so it is important to study the effect of climate in mixed stands with different silvicultures. In this work we present preliminary results of a thinning trial in a mixed stands of Pinus pinaster Ait. and Quercus pyrenaica Willd located in ?Montes de Toledo? (Central Spain). The main objectives of the experiment are to study the growth response of each species to thinning and to analyse if there are different climate-growth relationships between species and thinning treatments. The stand is a stratified mixture with a planted pine overstory and a natural regenerated oak lower stratum (understory). The experiment design is a Latin square of order three with the following treatments: heavy thinning from below (40% of basal area is removed), moderate thinning from below (25% of basal area is removed) and control or unthinned. The thinning was applied during the winter 2009-10. The position of all trees larger than 7.5 cm of diameter per plot was recorded. Diameter of all trees and heights of a sample were measured before the thinning. In each plot dendrometer bands were installed in 10 trees per species in order to study the effect of treatments on individual tree growth. Dedrometer bands were read every month to detect possible differences in growth pattern between treatments. These data were complemented with continued records registered with electronic dendrometer installed in one tree per species and treatment. Climatic variables and soil temperature and humidity were recorded with a weather station and soil sensors set up in the study site. Data from the first growing season after the thinning treatment (spring of 2010) will be presented in this work. The possible differences in total growth and in growth pattern between species and treatments will be analysed and discussed