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.

     

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.

     

Has global change induced divergent trends in radial growth of Pinus sylvestris and Pinus halepensis at their bioclimatic limit? The example of the Sainte-Baume forest (south-east France)

? It is agreed that climate (precipitation and temperature) influences the distribution of plant species. Near the margins of a species’ natural range, climate becomes limiting to physiological processes. There, climate change may be expected to have a significant impact on tree growth and the species’ ranges may be altered.

? In order to assess what influence climate change could exert on the distribution of pine species at their margin, radial growth trends in ring-width chronologies over the last century were analysed. In the French Mediterranean area where climate change is characterized by increased temperature, forest plots were selected along an altitudinal transect on the north-facing slope of the Sainte-Baume mountain (Bouches-du-Rhône, France) where the ranges of Pinus sylvestris and Pinus halepensis overlap.

? Two growth patterns were identified. For P. halepensis, radial growth has increased at all altitudes indicating that climate change has improved growth conditions of this species near the margin of its ecological range. For P. sylvestris, radial growth has increased only at low altitudes and even decreased at high altitudes.

? It must be deduced that the growth changes observed cannot be generalised either at the species level or at the geographical level and must be interpreted with great caution.

     

Productivity of temperate broad-leaved forest stands differing in tree species diversity

? Understanding the effects of tree species diversity on biomass and production of forests is fundamental for carbon sequestration strategies, particularly in the perspective of the current climate change. However, the diversity-productivity relationship in old-growth forests is not well understood.

? We quantified biomass and above-ground production in nine forest stands with increasing tree species diversity from monocultures of beech to stands consisting of up to five deciduous tree species (Fagus sylvatica, Fraxinus excelsior, Tilia spp., Carpinus betulus, Acer spp.) to examine (a) if mixed stands are more productive than monospecific stands, (b) how tree species differ in the productivity of stem wood, leaves and fruits, and (c) if beech productivity increases with tree diversity due to lower intraspecific competition and complementary resource use.

? Total above-ground biomass and wood production decreased with increasing tree species diversity. In Fagus and Fraxinus, the basal area-related wood productivity exceeded those of the co-occurring tree species, while Tilia had the highest leaf productivity. Fagus trees showed no elevated production per basal area in the mixed stands.

? We found no evidence of complementary resource use associated with biomass production. We conclude that above-ground productivity of old-growth temperate deciduous forests depend more on tree species-specific traits than on tree diversity itself.

     

Characteristics of a 20-year-old evergreen broad-leaved forest restocked by natural regeneration after clearcut-burning

? To elucidate the application of natural regeneration to the restocking of evergreen broad-leaved forests in the subtropics, the characteristics of a 20-year-old evergreen broad-leaved forest restocked by natural regeneration after clearcut-burning were studied in Okinawa, Japan. Within a 0.87 ha clearcut area with four 10 m × 10 m sampling plots, two burned and two unburned ones, the tree composition, tree species diversity and vegetation changes were investigated.

? The results showed that the species diversity, basal area and density of woody stems ? 1.0 m in height differed significantly among phanerophyte types, while they were not significantly different between the burned and unburned treatments. A vegetation census also revealed no obvious differences between the treatments.

? The primary dominant species, Castanopsis sieboldii, continued to dominate the secondary forest with a broad height distribution.

? The structural complexity and high tree species diversity of the regenerating forest after clearcut-burning provides no evidence of degeneration. We can predict that the regeneration forest may gradually develop into stands similar to pre-clearcut primary forest, and that natural regeneration may restore the high tree species diversity of the evergreen broad-leaved forests in Okinawa.

     

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.

     

Variations of plant and soil 87Sr/86Sr along the slope of a tropical inselberg

? From the summit downslope a granitic inselberg in French Guiana, soils and vegetation evolve from bare granite covered by cyanobacteria, to a savannah-type vegetation on thin patchy sandy accumulations, then to a low forest on shallow young soils and to a high forest on deep highly weathered ultisols.

? We have used element budgets and Sr isotopic variations in soils and plants to investigate the mineral nutrient supply sources of the different plant communities.

? Granite and atmospheric deposition have ⁸⁷Sr/⁸⁶Sr ratios of 1.3 and 0.71, respectively. The ⁸⁷Sr/⁸⁶Sr ratio of cyanobacteria (0.72) suggests granite weathering by cyanobacteria crusts. The ⁸⁷Sr/⁸⁶Sr ratio of the savannah-type vegetation is 0.73 and varies between 0.75 and 0.76 in the low and high forest leaf litter regardless of soil depth, age and degree of impoverishment.

? These almost constant ratio suggest that forest Sr comes from rainwater and from the summit of this inselberg, where it is released and redistributed along the slope, by surface flow, lateral redistribution of litter, and mineral particles. However, because of its very low content in the rock and soils, Ca is supplied to plants by atmospheric deposition.

     

Using cover measurements to estimate aboveground understorey biomass in Maritime pine stands

? Understorey plays a major role in forest fluxes and stocks balances, however this compartment is generally poorly quantified. Our objectives were to establish models to estimate understorey biomass using vegetation cover measurements and to investigate upscaling methodologies from stand to regional level.

? Understorey aboveground biomass measurements were undertaken in Maritime pine stands of mesohygric, mesic and dry moorlands in South West France.

? Average biomass stock in this compartment was estimated to 3.50 t DM ha^?1. The more abundant species groups varied with moorland types, with a higher relative contribution of herbaceous species (23.3%), bracken (59.2%) and mosses (31.6%) for mesohygric, mesic and dry moorlands, respectively. For each species group, we established significant relationships to estimate biomass using a volumetric index, based on cover and height measurements. No relationship between stand characteristics and understorey biomass was founded. We investigated the upscaling of these estimations to a several thousands hectare area using understorey cover measurements done along a regular spatial grid. The only significant correlation linked one satellite vegetation index to understorey biomass.

? We successfully developed empirical relationships to estimate the understorey biomass at the stand level. Further investigations could focus on the analysis of understorey variability over a finer space grid and the potential use of satellite vegetation indexes.

     

Long-term decrease of organic and inorganic nitrogen concentrations due to pine forest wildfire

? Growing concerns about fires and the increase of fire frequency and severity due to climate change have stimulated a large number of scientific papers about fire ecology. Most researchers have focused on the short-term effects of fire, and the knowledge about the long-term consequences of fires on ecosystem nutrient dynamics is still scarce.

? Our aim was to improve the existing knowledge about the long-term effects of wildfires on forestlabile N concentrations. We hypothesized that fires may cause an initial decline in organic and inorganic N availability, and in the amount of microbial biomass-N; this should be followed by the recovery of pre-fire N concentrations on a long-term basis. We selected a fire chronosequence in Pinus canariensis forests on La Palma Island (Canary Islands, Spain). These forests are under low anthropogenic atmospheric deposition, and forest management is completely lacking; wildfires are therefore the only significant disturbance. Soil samples were collected during the winter and spring at 22 burned and unburned plots.

? Fire produced a significant decrease in microbial biomass N, mineral N and dissolved organic N. Almost 20 y after fire, pre-fire levels of N concentrations had not recovered.

? These results demonstrate that P. canariensis forest soils have a lower resilience against fire than expected. The magnitude of these observed changes suggests that pine forest wildfires may induce long-term (2 decades) changes in soil and in plant primary production.

     

Interactive responses of Quercus suber L. seedlings to light and mild water stress: effects on morphology and gas exchange traits

? The combined effect of water stress and light on seedlings of forest species is a key factor to determine the best silvicultural and afforestation practices in the Mediterranean area.

? The aims of this work was (1) to determine the optimal light level for the early development of cork oak seedlings under mild water stress and (2) to test if the combined effect of water stress and light followed the trade-off, the facilitation or the orthogonal hypothesis.

? Shade reduced instantaneous photosynthetic rates and water use efficiency in cork oak. However, seedlings grown under moderate shade (15% of full sunlight) were capable to accumulate similar amount of biomass than those grown under more illuminated environments by increasing their specific leaf area. Absolute differences in net photosynthesis between light treatments were higher in well watered than in water stressed seedlings. However, the impact of both factors on overall growth was orthogonal.

? We concluded that cork oak development is impaired under deep shade (5% of full sunlight) but it can be optimal under moderate shade (15% of full sunlight) even under moderate water stress. Implications of these patterns on regeneration, cultivation and afforestation of cork oak are discussed.

     

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.

     

Wildfire changes the spatial pattern of soil nutrient availability in Pinus canariensis forests

? Soil resources are heterogeneously distributed in terrestrial plant communities. This heterogeneity is important because it determines the availability of local soil resources. A forest fire may change the spatial distribution of soil nutrients, affecting nutrition and survival of colonizing plants. However, specific information on the effects of ecosystem disturbance on the spatial distribution of soil resources is scarce.

? We hypothesized that, on a short-term basis, wildfire would change the spatial patterns of soil N and P availability. To test this hypothesis, we selected two Pinus canariensis forests burned in 2005 and 2000, respectively, and a third forest that was unburned since at least 1990 (unburned). We incubated ionic exchange membranes (IEMs) in replicated plots to estimate soil N and P availability and characterized the spatial pattern using SADIE (Spatial Analysis by Distance Indices).

? Mineral N, NO₃-N and PO₄-P availability, and aggregation and cluster indices for all nutrients were higher in the 2005 wildfire plots than in the 2000 wildfire and unburned plots.

? Our results suggest that surviving plants or new individuals becoming established in a burned area would find higher soil resources, but also higher small-scale heterogeneity in nutrients, which may have a major impact on the performance of individual plants and on the forest structure and dynamics.

     

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.

     

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).

     

Biomass component equations for Latin American species and groups of species

? Studies dealing with the estimation of biomass, site productivity and the contribution of forests to the global carbon balance require the use of allometric equations. There have been a great number of equations developed to estimate biomass components of trees and shrubs in various ecosystems. However, there are less literature compilations that address the calculations of biomass components.

? I report a total of 229 sets of allometric equations to estimate biomass components for 102 species in 72 different forest communities of arid, semi-arid, subtropical, tropical and temperate Latin-American ecosystems.

? The selection of the appropriate allometric model is a key element in the accurate estimation of biomass, stand productivity, carbon stocks and fluxes, and as a consequence, it is important to apply special effort to the selection and estimation of biomass equations.

? I also discuss statistical methods of parameter estimation and recommend the dissection of two conventional allometric equations when biomass studies are conducted on a wide range of diameters. In order to use nondestructive procedures of biomass estimation such as the fractal theory, the null hypothesis that the mean slope b value is equal to 2.67 was rejected for Latin American biomass species.

? This information is critical for the establishment of environmental projects that aim to estimate conventional parameters (i.e., productivity, habitat quality and fuel wood) as well as environmental features (i.e., stocks and fluxes of carbon and nitrogen).

     

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.

     

Temporal trends in the foliar nutritional status of the French, Walloon and Luxembourg broad-leaved plots of forest monitoring

? Two decades after the launching of the monitoring program of forest ecosystems in Europe (ICP forests), a unique data set is now available regarding the foliar nutritional status of the main broad-leaved species growing in much-diversified sites (soil, climate).

? This study focuses on the foliar concentration time series (1993 to 2005) of the French (RENECOFOR), Walloon and Luxemburg broad-leaved plots. The aim is to show long-term trends while taking the inter-plot and inter-annual variability into account.

? Two kinds of statistical processing were used to analyze the data on foliar chemistry: principal component analysis (PCAs) and linear mixed models. In general, the main temporal trends reveal a decrease in the foliar P concentration and a decrease in the foliar Ca concentration, except for European beech on acid soils.

? These temporal patterns confirm the trends previously observed and could be explained by the joint actions of several processes that influence tree nutrition in the long term: tree age, N and S deposition, harvesting and climate.

     

Integrating functional diversity into tropical forest plantation designs to study ecosystem processes

? The elucidation of relationships between biodiversity and ecosystem processes has been limited by the definition of metrics of biodiversity and their integration into experimental design. Functional trait screening can strengthen the performance of these designs.

? We suggest the use of Rao’s quadratic entropy to measure both functional diversity and phylogenetic diversity of species mixtures proposed for an experimental design, and demonstrate how they can provide complementary information.

? We also present an index assessing the statistical performance of these independent variables in different experimental designs. Measurement of independent variables as continuous vs. discrete variables reduces statistical performance, but improves the model by quantifying species differences masked by group assignments.

? To illustrate these advances, we present an example from a tropical forest tree community in which we screened 38 species for nine functional traits. The proposed TropiDEP design is based on the relative orthogonality of two multivariate trait axes defined using principal component analysis.

? We propose that independent variables describing functional diversity might be grouped to calculate independent variables describing suites of different traits with potentially different effects on particular ecosystem processes. In other systems these axes may differ from those reported here, yet the methods of analysis integrating functional and phylogenetic diversity into experimental design could be universal.

     

Effects of atmospheric and climate change at the timberline of the Central European Alps

? This review considers potential effects of atmospheric change and climate warming within the timberline ecotone of the Central European Alps. After focusing on the impacts of ozone (O₃) and rising atmospheric CO₂ concentration, effects of climate warming on the carbon and water balance of timberline trees and forests will be outlined towards conclusions about changes in tree growth and treeline dynamics.

? Presently, ambient ground-level O₃ concentrations do not exert crucial stress on adult conifers at the timberline of the Central European Alps. In response to elevated atmospheric CO₂ Larix decidua showed growth increase, whereas no such response was found in Pinus uncinata. Overall climate warming appears as the factor responsible for the observed growth stimulation of timberline trees.

? Increased seedling re-establishment in the Central European Alps however, resulted from invasion into potential habitats rather than upward migration due to climate change, although seedlings will only reach tree size upon successful coupling with the atmosphere and thus loosing the beneficial microclimate of low stature vegetation.

? In conclusion, future climate extremes are more likely than the gradual temperature increase to control treeline dynamics in the Central European Alps.

     

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.