Disentangling the effect of size and competition: a growth model for Aucoumea klaineana

• Context Okoume (Aucoumea klaineana) is the most important timber species in Gabon, while being classified as vulnerable in the IUCN red list. Thousands of growth measurements for this species have been performed since the 1950s but, because of a lack of integrated analysis, did not bring a consistent view on its growth pattern.
• Aims This study aims at disentangling the effects of tree size and local competition on tree growth.
• Methods A growth model was fitted for okoume, using data from seven sites in Gabon and Congo. The growth model was designed to be useful for forest management, which means that it relied on variables that could be computed using forest inventory data.
• Results A lognormal growth model with a negative response to stand basal area was selected. A significant residual site effect on growth was found, with a slower growth in the sites near the border of the natural range of okoume than in the sites far from it.
• Conclusion Growth strongly responded to local stand attributes such as tree density and basal area. Growth decrease with stand age in monodominant okoume stands was correctly predicted, although tree age was not incorporated into the model.

     

Growth response of Pinus pinaster Ait. to climatic variables in central Spanish forests

? The effect of climatic variables (temperature and precipitation) on radial growth of the Mediterranean Maritime pine (Pinus pinaster Ait.) was studied using dendrochronological techniques in the Iberian Peninsula.

? Ten tree-ring width chronologies, along the central distribution area of the species, were built. Chronology variability was analysed using Principal Component Analysis (PCA) for the period 1952–2005.

? The first principal component (PC1) explained 56% of tree-growth variability. Tree-growth association with climate was analysed at regional and local scales using correlation coefficient and bootstrapped response functions.

? Radial growth at both scales was positively correlated with rainfall during and prior to the growing season at all sites, and with summer rainfall before the growing season at five sites. Mean temperature effect changed according to the sampling site, from non-significant at the highest sites to significant (positive relationship in winter) at the lowest sites. Growth season temperature also had a negative effect.

? The Kalman filter was applied to estimate changing association between growth and climate over-time. Results suggested a change in association, initiated in the 80s, from non-significant to significant (*p < 0.05) at six of the sampling sites.

? Pinus pinaster is an accurate species for analysing tree-growth association with climate and for studying plant behaviour under global change conditions.

     

Black pine (Pinus nigra Arn.) growth divergence along a latitudinal gradient in Western Mediterranean mountains

? Most studies of tree-growth and climate report positive responses to global warming in high latitudes and negative responses at lower ones.

? We analyzed tree-ring width of Pinus nigra Arn. along a 500 km latitudinal transect in the Iberian Peninsula to study the temporal trend and climate forcing in tree radial growth during the last century.

? Tree growth was enhanced by cool summers and moist cold seasons. Increased moisture stress has decreased tree growth rates. However, we present evidence of growth increases in some trees in all sampled populations after 1980’s. Climate change negatively (positively) affected between 72% (5%) of trees in the southern populations and 40% (25%) in the north Trees with positive growth trends were favored by winter temperatures and their abundance was inversely correlated with forest productivity.

? Our findings add evidences of tree growth divergence in the Mediterranean basin and show the gradual transition between forests where positive (temperate and boreal) and negative (Mediterranean) growth trends dominate.

     

A merchantable volume system for Pinus sylvestris L. in the major mountain ranges of Spain

? Accurate estimation of stem volume makes it possible to estimate the monetary value of one of the many commodities and services that forests provide to society, i.e. timber.

? In the present study a compatible volume system for Scots pine in the major mountain ranges of Spain (the Pyrenees, Southern Iberian Range, Northern Iberian Range, Soria and Burgos Mountains, Central Range and Galician Mountains) was developed with data from 2 682 destructively sampled trees. Several well-known taper functions were evaluated. A second-order continuous-time autoregressive error structure was used to correct the inherent autocorrelation in the hierarchical data, allowing the model to be applied to irregularly spaced and unbalanced data.

? The compatible segmented model of Fang et al. (2000) best described the experimental data. It is therefore recommended for estimating diameter at a specific height, height to a specific diameter, merchantable volume, and total volume for the six mountain ranges analyzed.

? The non-linear extra sum of squares method indicated differences in mountain range-specific taper functions. A different taper function should therefore be used for each mountain range in Spain.

     

A model bridging distance-dependent and distance-independent tree models to simulate the growth of mixed forests

? It is widely believed that distance-independent tree models fail to take into account the complexity of mixed stands due to the fact that spatial structure often has a greater impact on growth and dynamics in mixed stands than in pure stands. On the other hand, distance-dependent tree models are difficult to use because they require a map of the stand, which is not only very costly but also impracticable in a routine management context.

? This paper reports the development of a model bridging distance-dependent and distanceindependent tree models, and that is designed to simulate the growth of a mixed forest. The model used distributions of the number of neighbours to reconstruct tree neighbourhoods and compute the competition indices needed as inputs to the growth model.

? Data were collected from a mixed forest of sessile oak and Scots pine in central France. The study showed that local competition indices explained a significant proportion of growth variability and that intraspecific competition was greater than interspecific competition. The model based on neighbourhood distributions gave consistent predictions compared to a distance-dependent model.

? This type of model could be used instead of distance-dependent models in management contexts.

     

Site preparation and competing vegetation control affect loblolly pine long-term productivity in the southern Piedmont/Upper Coastal Plain of the United States

? A site preparation study was established in 1986 to evaluate the effect of different site preparation treatments on growth and yield of loblolly pine (Pinus taeda L.) plantations on the Piedmont and Upper Coastal Plain regions of the southern United States. Site preparation treatments included: (1) burn only, (2) chop-burn, (3) shear-pile-disk, (4) chop-herbicide-burn, (5) herbicide-burn, and (6) herbicide-burn-herbicide.

? The data from the available 19 installations at age 21 were analyzed with separate analysis of variance and a multilevel nonlinear mixed-effects modeling approach.

? The herbicide-burn-herbicide treatment significantly increased average Dbh, height, basal area and volume per hectare compared to all other treatments. The burn only treatment consistently ranked worst and was followed by the chop-burn treatment. The shear-pile-disk and chop-herbicide-burn treatments had similar overall growth pattern, and will approach the same level of pine volume as the herbicide-burn treatment.

? Loblolly pine mean annual increment in volume (m³ ha^?1 y^?1) at age 21 by treatment were: herbicide-burn-herbicide (17.9), shear-pile-disk (16.1), herbicide-burn (15.9), chop-herbicide-burn (15.4), chop-burn (14.3), and burn (11.2).

? An additional chop or herbicide treatment to the burn treatment significantly increased loblolly pine yield. Complete control of both herbaceous and woody completion enhanced long-term pine productivity.

     

Fuel and stand conditions in an isolated, unmanaged forest landscape in central Oregon

? Mature, unmanaged forests in western North America provide important reference conditions for managers, yet little is known about fuel composition and the factors that influence fuel accumulation in such stands. Our objectives were to characterize fuels in a passively managed landscape of dry forests in central Oregon and identify environmental factors influencing fuel accumulation.

? Ordination techniques and analysis of variance revealed no statistical differences in total fuel loads across a wide range of environmental conditions.

? Individual fuel size classes, however, did vary by stand location and composition. Interior stands had more 1- to 100-h fuels and snags than stands at or near the edge, stands dominated by ponderosa pine had fewer small-diameter fuels and snags, lodgepole pine stands had more 1-h fuels and snags, and white fir stands had more 10-h fuels, duff, and snag basal area.

? Tree species, density and age, and years since last fire were the most important environmental variables, explaining 23% of the variation in fuels.

? Our results present further evidence that fuel composition is highly variable at many spatial and temporal scales. They also provide useful baseline information for managers of fire-prone western forests.

     

Rank reversals in tree growth along tree size, competition and climatic gradients for four forest canopy dominant species in Central Spain

? Interspecific differences in tree growth patterns with respect to biotic and abiotic factors are key for understanding forest structure and dynamics, and predicting potential changes under climate change.

? Repeated observations from the Spanish Forest Inventory (SFI) were used to parameterize maximum likelihood estimators of tree growth as a function of tree size, competition indices and climate for Pinus pinaster, P. sylvestris, Quercus ilex and Q. pyrenaica.

? Significant responses to both biotic and abiotic factors were found, with interspecific differences in species performance along competition, temperature and precipitation gradients. Q. ilex was the species most tolerant to competition while P. pinaster was the species most sensitive to climatic variation. Species relative positions shifted along gradients of these factors with rank reversals in species performance along size, competition and climatic gradients.

? The results based on average growth matched previous forestry classifications and experimental studies on relative growth rate (RGR).

? When examining growth along studied abiotic and biotic gradients, a mismatch was found between species performance ranks as predicted by our models and information derived from previous knowledge. Those discrepancies highlight the relevance of ontogeny and environmental heterogeneity in defining species performance along competition gradients.

     

Potential productivity of forested areas based on a biophysical model. A case study of a mountainous region in northern Spain

? Today’s forest managers face a number of important challenges involving an increasing need for precise estimates of forest structure and biomass, potential productivity or forest growth. The objective is to develop a model for potential productivity in a mountainous region of Spain. The model combines climatic, topographic and lithological data using a variant of a traditional biophysical model: the Paterson index.

? In a first approach, the climatic productivity is assessed by modelling the required parameters using different geostatistical techniques and software supported by GIS. A second approach includes the correction of the former productivity classes considering the different lithological facies. The potential forest productivity model involves the integration of both models.

? Finally, data from the National Forest Inventory (NFI) are used to compare the real and potential yield data within different regions of the studied area.

? The results of these analyses demonstrate the usefulness of the model, particularly in mountainous regions, where no significant differences are found between the data from the NFI and the model, but they also show the discrepancies between the estimates and real data when the latter are considered for different tree species, diameter classes or management.

     

Is forest mushroom productivity driven by tree growth? Results from a thinning experiment

? Most of the edible forest mushrooms are mycorrhizal and depend on carbohydrates produced by the associated trees. Fruiting patterns of these fungi are not yet fully understood since climatic factors alone do not completely explain mushroom occurrence.

? The objective of this study was to retrospectively find out if changing tree growth following an increment thinning has influenced the diversity patterns and productivity of associated forest mushrooms in the fungus reserve La Chanéaz, Switzerland.

? The results reveal a clear temporal relationship between the thinning, the growth reaction of trees and the reaction of the fungal community, especially for the ectomycorrhizal species. The tree-ring width of the formerly suppressed beech trees and the fruit body number increased after thinning, leading to a significantly positive correlation between fruit body numbers and tree-ring width.

? Fruit body production was influenced by previous annual tree growth, the best accordance was found between fruit body production and the tree-ring width two years previously.

? The results support the hypothesis that ectomycorrhizal fruit body production must be linked with the growth of the associated host trees. Moreover, the findings indicate the importance of including mycorrhizal fungi as important players when discussing a tree as a carbon source or sink.

     

Influence of several tree traits on rainfall partitioning in temperate and boreal forests: a review

? In forests, rainfall is partitioned into intercepted water (IW), throughfall (TF) and stemflow (SF). We reviewed the majority of published works comparing water flows among tree species in temperate and boreal forests to test the effect of several tree traits on water flows.

? We hypothesized that water flows differed between evergreen and deciduous species, and according to successional status and bark roughness. We also investigated that water flows would be explained by stand variables such as basal area.

? Linear mixed models fitted on reviewed data showed that evergreens had a lower TF than deciduous trees (decrease of 13.9% of total precipitation year-round and 8.4% over the growing period). Similar results were found for conifers compared to broadleaves. TF also declined with the successional status from pioneer to late-successional tree species. SF decreased with bark roughness from smoother to rougher bark. Evergreens had water flows that were dependent on age of the stand, especially for TF which increased by 15.6% of total precipitation from young to adult forests.

? The large scale of TF differences between tree genera together with specific transpiration amounts and rooting features highlighted in other studies should result in significant differences in soil water content among tree species. This may have consequences on stand fitness and growth, and understory vegetation.

     

Modelling ozone fluxes to forests for risk assessment: status and prospects

? Risk assessment of ozone effects on forests is gradually moving from concentration-based exposure metrics to a more complicated approach that requires modelling of ozone fluxes to trees.

? This study reviews the status of the DO₃SE stomatal flux model employed within the Convention on Long-range Transboundary Air Pollution, describing a range of applications and identifying major research needs, especially in the context of support that could be provided by the International Co-operative Programme on Assessment and Monitoring of Air Pollution Effects on Forests.

? The most urgent development need for DO₃SE is the modelling of the soil moisture status and its effect on stomatal conductance. Furthermore, the data related to the physical characteristics and the seasonal dynamics of physiological activity of vegetation continue to pose problems.

? There is a clear need for more extensive validation of models and risk estimates using more rigorous statistical procedures and comparisons with flux networks and satellites.

? The current large-scale forest monitoring activities provide only limited possibilities for flux modelling, but could be enhanced by introducing a new monitoring strategy outlined here.

     

A generic model of thinning and stand density effects on forest growth, mortality and net increment

? For assessing forest thinning effects at large (i.e. continental) scale, data scarcity and technical limitations prevent the application of localized or individual-based thinning models.

? Here we present a simple general framework to analyze and predict the effects of thinning on growth and mortality, including the following stand density development. The effects are modeled in relative terms so that the model can be parameterized based on any thinning experiment that includes an unthinned control, regardless of site conditions and stand age.

? The model was tested against observed thinning effects on growth and mortality from five temperate and boreal species (all species pooled r ² = 0.51). It predicted a maximum increase in net stem biomass increment of 16% and a reduction in density-related mortality of 75% compared to un-thinned conditions at stand densities of around 70% of the maximum (increment optimal density).

? A sensitivity analysis revealed overlapping ranges of near optimal density (net increment within 95% of optimal) among all tested species, suggesting that one thinning scenario can be used for many species. The simple and general formulation of thinning effects based on only five parameters allows easy integration with a wide range of generic forest growth models.

     

A new bioclimatic model calibrated with vegetation for Mediterranean forest areas

? Water availability is one of the main factors explaining flora composition and growth in Mediterranean regions, where it may decline with climate change.

? Our goal was to develop a model for forest site assessment in Mediterranean environments, focusing on water availability to assess potential vegetation composition and productivity in any places, whatever their level of disturbance.

? We designed a statistical model, using global climatic and geographic variables, as well as detailed local topographic and edaphic variables, to compute a bioclimatic index for Mediterranean forest environments. This model was calibrated in France with a flora index from 325 old forests. The model explained 80.3% of the flora index variance. The method fills a gap in existing models, bridging scales from the region to forest sites.

? Beyond its theoretical aspect, it was designed to allow practical tools to be derived from it for decision-making and management, such as the assessment of climate change impact on vegetation, and of forest productivity. Its development and adaptation is possible in other Mediterranean regions, and in any region where water is one of the main limiting factors.

     

Modelling site quality and individual-tree growth in pure and mixed Pinus brutia stands in north-east Greece

? Simulation tools, based on individual tree growth and mortality models can produce the most detailed predictions of forest stand development under different management schedules. These models allow the manager to predict the development of any type of stand (even- and uneven-aged, and pure and mixed stands).

? Different model approaches and predictors are required for pure even-aged or mixed uneven-aged forest stands. This study developed and compared two sets of models which enable tree-level simulation of the development of pure and mixed stands of Pinus brutia in north-east Greece. The first set of models for even-aged forestry consists of site index models, diameter growth models, tree height models, and mortality models. The second set, which is for uneven-aged forestry, uses a past growth index instead of a site index.

? The simulations and overall fitting statistics suggest that the two types of models provide realistic and accurate predictions of forest stand development and allow one to simulate the development of complex Pinus brutia stand structures in Dadia National Park forests.

? The advantages of the two approaches are discussed and it is suggested that the growth index is an effective predictor of site quality and the set of models which used such variable as predictor performed in a similar way as the models using site index, which require more information and a given stand structure (even-aged).

     

Oak somatic and gametic embryos maturation is affected by charcoal and specific aminoacids mixture

? Development of both somatic and gametic embryogenesis has many applications in clonal forestry and genetic improvement, for instance as mass-propagation of genetically improved plants and production of pure lines through doubled-haploid plant regeneration from gametic embryos.

? The goal of this work was to improve growth, maturation and plantlet regeneration of cork oak (Quercus suber L.) embryos from both somatic and gametic origin.

? Activated charcoal promoted a significant increase in growth in terms of relative size and weight of both somatic and doubled-haploid embryos, as well as a more efficient control of secondary somatic embryogenesis during development. A significant interaction was also observed with amino acid nutrition. While some amino acids (i.e., glutamine, arginine or asparagine) did not show significant differences with the controls, a mixture of these three amino acids or gamma amino butyric acid stimulated embryo growth. The highest survival rate during acclimation of plantlets from both somatic and doubled-haploid origin was obtained when the embryos had been previously cultured on basal medium with 3% sucrose and 1% activated charcoal for two months at 4 °C and germinated on medium supplemented with 6-benzylaminopurine and indole-3-butyric acid.

? We obtained more than 900 cork oak plantlets acclimated from several embryogenic lines, with a high survival rate, demonstrating that this methodology is applicable for large scale plantlet production. We also report the first regeneration of doubled-haploid plantlets in cork oak.

     

Comparing volume growth in pure and mixed stands of Pinus sylvestris and Quercus pyrenaica

? 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.

? 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.

     

Diachronic analysis of individual-tree mortality in a Norway spruce stand in the eastern Italian Alps

? Understanding tree mortality processes across time requires long term studies. Spatiotemporal patterns of mortality in a 200 years-old mono-layered Norway spruce stand were evaluated to determine what factors affected individual-tree mortality.

? We performed an analysis on two surveys (1993 and 2005) in a 1-ha permanent plot in the Paneveggio forest (Eastern Italian Alps). Tree diameter and age distribution between surveys were compared. We examined spatial patterns of living and dead trees before 1993, in 1993 and in 2005 using univariate and bivariate Ripley’s K(d) function, and a kernel estimator of local crowding. A logistic model was used to assess the effects of diameter, age, recent growth and competitive pressure on tree mortality.

? Spatial pattern analysis indicated mortality was associated to tree neighbourhood (neighbour effect at 2–5 m). An increment of regularization of tree spatial pattern occurred due to density-dependent mortality. Logistic regression showed tree diameter and recent growth were determinant on mortality risk during the monitoring period.

? Even if the stand is relatively aged, mortality dynamics are those typical of stem exclusion stage. Mortality was related to competitive dynamics, and small suppressed trees with slow growth rate had higher probability to die.

     

Fecundity and larval development of Monochamus galloprovincialis (Coleoptera Cerambycidae) in experimental breeding

?The study aimed at clarifying basic life traits (fecundity, mortality and development) of Monochamus galloprovincialis, the vector of Bursaphelenchus xylophilus, the causative agent of the Pine Wilt Disease, in Portugal.

? Mating and oviposition experiments were conducted on P. sylvestris logs under laboratory conditions during two years. Larval development was followed outdoors.

? Fecundity was high, with 138.2 eggs per female.

? For the first time, the number of larval instars (4) in M. galloprovincialis was recognized and instars were described.

? Informations are provided on the shape and size of the larval galleries. Boring of galleries in sapwood and heartwood started at the third instar. All instars could overwinter but adults emerged simultaneously resulting in a univoltine life cycle for the majority of the individuals.

? An obligatory diapause in the forth instar was assumed. However, 8.1% of the insects had a two year development.

? Important differences in mortality were noted between the two experimental years, which could be explained by differences on the size of the breeding logs.

? There is a high biological proximity between M. galloprovincialis populations in France and those in Portugal and Northern Europe. However, a higher fecundity and longevity was observed in France, as well as a higher percentage of insects with a two year development, compared with portuguese populations.

     

Compared effects of competition by grasses (Graminoids) and broom (Cytisus scoparius) on growth and functional traits of beech saplings (Fagus sylvatica)

? The effect of two weeds (grasses and broom, Cytisus scoparius) competition on the growth and functional traits of European beech saplings (Fagus sylvatica) was investigated in an experimental plantation in the French Massif central.

? We hypothesized that grasses would have a much more harmful effect than broom on beech growth through strong competition for soil water and nitrogen.

? A randomized block design was used with three separate blocks, each possessing three types of vegetation; grasses (mainly Agrostis capillaris, Holcus mollis and H. lanatus), broom, and bare soil as a control.

? Two years after plantation beech sapling growth was significantly greater in bare soil than with competition from grasses, and was intermediate on broom. Growth was positively correlated with beech leaf nitrogen content (N_m) and xylem water potential (Ψ_x). N and Ψ_x were dependent, with nitrogen uptake probably decreasing with increasing drought. There was no clear correlation between growth and light transmitted 10 cm below the apex of the saplings.

? Beech leaf mass on an area basis (LMA) was correlated with the three resource variables (water, nitrogen and light) and with growth, confirming that LMA is a sensitive functional trait integrating the degree of stress experienced by the saplings.

? The results suggest that broom is less competitive than grasses and, if properly managed on a sufficiently well-watered site, can generate satisfactory conditions for beech establishment and growth.