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.

     

A comprehensive framework of forest stand property–density relationships: perspectives for plant population ecology and forest management

? Context

There are many stand property–density relationships in ecology which represent emergent properties of plant populations. Examples include self-thinning, competition–density effect, constant final yield, and age-related decline in stand growth. We suggest that these relationships are different aspects of a general framework of stand property–density relationships.

? Aims

We aim to illustrate the generalities and ecological implications of stand property–density relationships, and organize them in a comprehensive framework.

? Methods

We illustrate relationships between stand property and density (1) at one point in time, (2) over time, and (3) independent of time. We review the consequences of considering different variables to characterize stand property (mean tree size, mean tree growth, stand growth, stand yield, stand leaf area).

? Results

We provide a framework that integrates the broad categories of stand property–density relationships and individual expressions of these relationships. For example, we conclude that constant final yield is a special case of the growth–growing stock relationship for life forms were yield is a reasonable approximation of growth (non-woody plants).

? Conclusion

There is support in the literature for leaf area being broadly integrative with respect to various expressions of stand property–density relationships. We show how this is and suggest implications for plant population ecology and forest management.     

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.

     

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).

     

Models of the longitudinal distribution of ring area as a function of tree and stand attributes for four major Canadian conifers

Context

It is widely accepted that ring area increment generally increases from the tree apex to the crown base and is more-or-less constant below the crown base (Pressler’s law), but few quantitative models of this distribution have been developed.

Aims

The aim of this study was to develop a model of ring area increment using easily obtained crown features and other tree or stand characteristics in order to further the understanding and prediction of tree growth, form, and wood quality.

Methods

The models were fit to stem analysis observations from white spruce, black spruce, balsam fir, and lodgepole pine.

Results

In the final model, which includes tree crown and stand variables, ring area increment within the crown region was slightly curvilinear, the slope of ring area increment below the crown was non-zero, and the effect of butt swell was appreciable up-to a relative height of 0.10.

Conclusions

The high accuracy of the mixed effects model suggests that the three-component model form is appropriate for describing ring area profiles, whereas some tree-to-tree variation remains unexplained. The tree and stand variables used in these models can be easily measured in the field or obtained from remote sensing techniques.     

Properties of boundary-line release criteria in North American tree species

? Boundary line release criteria are increasingly applied to evaluate forest disturbance histories from tree-ring data. However, a number of important properties central to the technique have not been evaluated, including: (i) the ability of boundary line release criteria to standardize releases across various sites, species, and tree life stages (ii) the minimum sample sizes necessary for developing boundary lines, and (iii) the degree to which the criteria can resolve the degree of crown exposure following a disturbance event.

? In an analysis of eleven North American tree species, boundary line release criteria do not fully compensate for declines in release response a tree experiences with increasing age and size, with the exception Tsuga canadensis.

? A bootstrapping analysis indicates that approximately 50 000 ring width measurements are necessary to develop boundary line release criteria for a given species.

? In a Quercus prinus stand, boundary line release criteria better predict the degree of crown exposure following a disturbance than an earlier running mean technique.

? Despite certain limitations, boundary line release criteria have the potential to standardize release calculation across most life stages of a tree, and possibly among sites and species.

     

Ten years of fluxes and stand growth in a young beech forest at Hesse, North-eastern France

? Water and carbon fluxes, as measured by eddy covariance, climate, soil water content, leaf area index, tree biomass, biomass increment (BI), litter fall and mortality were monitored for 10 successive years in a young beech stand in Hesse forest (north-eastern France) under contrasting climatic and management conditions.

? Large year-to-year variability of net carbon fluxes (NEE) and to a lesser extent, of tree growth was observed. The variability in NEE (coefficient of variation, CV = 44%) was related to both gross primary production (GPP) and to variations in total ecosystem respiration (TER), each term showing similar and lower interannual variability (CV = 14%) than NEE. Variation in the annual GPP was related to: (i) the water deficit duration and intensity cumulated over the growing season, and (ii) the growing season length, i.e. the period of carbon uptake by the stand. Two thinnings occurring during the observation period did not provoke a reduction in either GPP, water fluxes, or in tree growth. Interannual variation of TER could not be explained by any annual climatic variables, or LAI, and only water deficit duration showed a poor correlation. Annual biomass increment was well correlated to water shortage duration and was significantly influenced by drought in the previous year.

? The relationship between annual NEE and biomass increment (BI) was poor: in some years, the annual carbon uptake was much higher and in others much lower than tree growth. However this relationship was much stronger and linear (r ² = 0.93) on a weekly to monthly time-scale from budburst to the date of radial growth cessation, indicating a strong link between net carbon uptake and tree growth, while carbon losses by respiration occurring after this date upset this relationship.

? Despite the lack of correlation between annual data, the NEE and BI cumulated over the 10 years of observations were very close.

? On the annual time-scale, net primary productivity calculated from eddy fluxes and from biological measurements showed a good correlation.

     

Climate-growth variability in Quercus ilex L. west Iberian open woodlands of different stand density

? We present the longest tree-ring chronology (141 y) of Quercus ilex L. (holm oak), and discuss the species climate-growth relationships and the influence of stand density on tree sensitivity to climate.

? Similarly to Quercus suber L., the most influential climatic variables upon holm oak growth were late spring and early summer precipitation, which enhanced growth, and high temperatures in the previous August and current July, which negatively affected growth.

? High density stands responded to similar climatic factors as low density stands, but their response was generally weaker. Holm oak sensitivity to climate has increased in recent decades, which might be related to increasing temperatures in the region. Sensitivity was higher in low density stands. Additionally, the effect of summer stress on growth seems to have increased during the same period, similarly to other species in the Iberian Peninsula, suggesting that trees are more vulnerable to climatic changes.

? Stand density could buffer the response to climate by smoothing climatic extremes. Nevertheless, the effect of competition might reverse this positive effect at the individual tree level. Precautions should be taken before providing management guidelines regarding the effect of climate change and stand density on holm oak.

     

Which factors most influence heartwood distribution and radial growth in plantation teak?

? To investigate the effect of climate on radial growth in young plantation grown teak (Tectona grandis L.), growth ring width was measured in 105 trees and correlated to precipitation and temperature data.

? The social status of trees within the stand was also determined and cross-sectional area (CSA) for the trunk correlated to the proportion of heartwood (HW) within the tree. HW develops asymmetrically in leaning stems of some conifer species, but it is not known if this phenomenon also occurs in broadleaf species. Therefore, we measured HW proportion in leaning and straight stems, along with the number of growth rings in the HW.

? Annual ring width depended strongly on mean monthly temperature during the rainy season and the most significant relationships were found corresponding to the months of June and July. With regard to the weaker relationship between precipitation and radial growth, correlations were highest during the period of bud-break at the beginning of the rainy season.

? The very high stand density affected radial growth, particularly in suppressed trees, which responded little to thinning operations. HW formation was greatest in dominant trees, and was highly regressed with stem CSA.

? Therefore, rapid growth of young stands should be encouraged by reducing stand density. Asymmetric HW formation occurred in both leaning and straight trees, and was significantly greater along the upper sides of leaning stems. It is probable that this eccentric HW formation is linked to mechanical loading on the tree.

     

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.

     

Thinning increases cone production of stone pine (Pinus pinea L.) stands in the Northern Plateau (Spain)

Context

Edible stone pine (Pinus pinea L.) nut is a forest product which provides the highest incomes to the owners of stone pine forests.

Aim

The objective of this work is to evaluate the effect of first thinning on growth and cone production in an artificially regenerated stand in order to determine optimum intensity.

Methods

A thinning trial was installed in 2004 to compare two thinning regimes (heavy and moderate) and a control treatment. From 2004 to 2012, six inventories of forest attributes were carried out, and the cone crop was harvested annually. We evaluated the effect of thinnings on growth using repeated measures analysis of variance with a mixed model approach. With regards to cone production, we first estimated the probability of finding cones in a tree by applying a generalized mixed model and then estimated cone production by using a mixed model, including climatic variables.

Results

We found that thinning had a positive influence on tree diameter increment. Thinning increased the probability of finding cones and cone production. However, significant differences between heavy and moderate thinnings were not found.

Conclusion

We recommend early silvicultural treatments in stone pine stands to favor the development of trees and larger edible pine nut production.     

Ecology of Armillaria species on silver fir (Abies alba) in the Spanish Pyrenees

? We describe the distribution and the ecology of three Armillaria species observed in silver fir (Abies alba) forests of the Pyrenees.

? We surveyed the presence and abundance of Armillaria above and belowground in 29 stands. Isolates were identified by the PCR-RFLP pattern of the IGS-1 region of their ribosomal DNA. We measured several ecological and management parameters of each stand in order to describe Armillaria infected sites.

? Armillaria cepistipes was the most abundant of three species observed. Armillaria gallica was dominant in soils with a higher pH and at lower elevations. Armillaria ostoyae seemed to be more frequent in stands where A. alba recently increased its dominance relative to other forest tree species. Thinning activities correlated with an increased abundance of Armillaria belowground. In 83% of the stands the same Armillaria species was observed above and belowground.

? It seems that in a conifer forest, A. cepistipes can be more frequent than A. ostoyae, a virulent conifer pathogen. Since logging is related to a higher abundance of Armillaria in the soil, the particular Armillaria species present in a given stand could be considered an additional site factor when making management decisions.

     

Development of alder carr after the abandonment of wet grasslands during the last 70 years

? The secondary succession of wet grasslands to communities of alder carr dominated by Alnus glutinosa was recorded in different parts of Europe during the 20th century. However, knowledge of such development of alder carr remains insufficient.

? The development of alder carr was reconstructed at five sites in the Czech Republic, using historical aerial photographs and methods of dendrochronology. The aims were to investigate the succession from wet grasslands to alder carr at sites previously intensively managed for agriculture and to find out the role of fluctuations in the groundwater table, caused by artificial drainage channels, in the observed stand dynamics and tree growth.

? The spread of forest (i.e., an increase in forest cover) predominated until the 1970s at all sites. This trend was disrupted by a large-scale dieback of forest stands in four of the five sites after the 1970s, followed by an increase in patch heterogeneity, as indicated by landscape metrics. The radial growth increment in Alnus glutinosa has been affected predominately by local environmental factors, probably including the changing degree of waterlogging. Forest dieback was presumably connected with a lesser extent of drainage channels.

? Our results indicate that observed successional pathways at sites of alder carr were probably caused by local changes in the groundwater table.

     

Tree diameter, height and stocking in even-aged forests

? Empirical observations suggest that in pure even-aged forests, the mean diameter of forest trees (D, diameter at breast height, 1.3 m above ground) tends to remain a constant proportion of stand height (H, average height of the largest trees in a stand) divided by the logarithm of stand density (N, number of trees per hectare): D = β(H ? 1.3)/ ln(N).

? Thinning causes a relatively small and temporary change in the slope β, the magnitude and duration of which depends on the nature of the thinning.

? This relationship may provide a robust predictor of growth in situations where scarce data and resources preclude more sophisticated modelling approaches.

     

Growth response to climate and drought change along an aridity gradient in the southernmost Pinus nigra relict forests

Context

Tree populations at the rear edge of species distribution are sensitive to climate stress and drought. However, growth responses of these tree populations to those stressors may vary along climatic gradients.

Aims

To analyze growth responses to climate and drought using dendrochronology in rear-edge Pinus nigra populations located along an aridity gradient.

Methods

Tree-ring width chronologies were built for the twentieth century and related to monthly climatic variables, a drought index (Standardized Precipitation–Evapotranspiration Index), and two atmospheric circulation patterns (North Atlantic and Western Mediterranean Oscillations).

Results

Growth was enhanced by wet and cold previous autumns and warm late winters before tree-ring formation. The influence of the previous year conditions on growth increased during the past century. Growth was significantly related to North Atlantic and Western Mediterranean Oscillations in two out of five sites. The strongest responses of growth to the drought index were observed in the most xeric sites.

Conclusion

Dry conditions before tree-ring formation constrain growth in rear-edge P. nigra populations. The comparisons of climate-growth responses along aridity gradients allow characterizing the sensitivity of relict stands to climate warming.     

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.

     

Estimating growth in beech forests: a study based on long term experiments in Switzerland

? This contribution presents a dynamic stand growth model for Beech (Fagus sylvatica L.) forests, based on a dataset provided by the Swiss Federal Institute for Forest, Snow and Landscape Research WSL, Birmensdorf. The dataset includes 143 research plots, covering a wide range of growing sites and providing up to 16 interval measurements per research plot.

? The objective of this research is to complement the range of existing beech growth models by bridging the gap between the historical yield tables and the single tree growth models. The specific aim is to develop transition functions which will project three state variables (dominant height, basal area and number of trees per hectare) at any particular time, in response to any arbitrary silvicultural treatment.

? Two of the transition functions were derived using the generalized algebraic difference approach (GADA), the third one was derived with the algebraic difference approach (ADA). All the functions were fitted simultaneously using iterative seemingly unrelated regression and a base-age-invariant method. The influence of thinnings on basal area growth was included by fitting different transition functions for thinned and unthinned stands.

? The overall model provides satisfactory predictions for time intervals up to 20 years. The new model is robust and its relatively simple structure makes it suitable for economic analysis and decision support.

     

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 annual weather on growth of pedunculate oak in southern Sweden

? A network of oak (Quercus robur L.) chronologies containing 49 sites and 635 single trees was analysed to identify weather variables affecting annual tree-ring increment dynamics in southern Sweden during 1860–2000.

? We analysed (1) the growth response of oak to non-extreme weather, and (2) the temporal and spatial patterns of regional growth anomalies (pointer years) and associated climatic extremes resolved on a monthly scale.

? Growth was controlled by precipitation in the current (June–July) and the previous growing season (August) in 48% and 22% of all sites, respectively. Temperature during July of the current year and August of the previous year was negatively correlated with growth in 29% and 43% of the sites, respectively. Growth was positively correlated with temperature in October of the previous season in 72% of the sites. The most extensive growth anomaly occurred in 1965 and was probably caused by intrusion of cold Arctic air masses into the region at the end of March that year.

? During climatically non-extreme years, oak growth is driven mostly by the dynamics of summer precipitation. Many of the negative growth anomalies, however, were associated with temperature extremes. Southern Swedish oak pointer years tend not to coincide with the pan-European oak pointer years.