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).

     

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.

     

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.

     

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.

     

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.

     

A tree distance-dependent growth and yield model for naturally regenerated pure uneven-aged maritime pine stands in central inland of Portugal

? Context

Maritime pine (Pinus pinaster Aiton) is one of the most important Portuguese species, growing in pure stands ranging from even-aged to multi-aged structures. Current growth and yield models were developed only for even-aged, managed stands and/or for very specific regions of Portugal.

? Aims

This paper focuses on the validation of the existing size-class model PBRAVO, adapted to even-aged stands, and on the subsequent development of a single tree distance-dependent growth and yield model (PBIRROL), both in distance-independent and distance-dependent versions, for uneven-aged stands.

? Methods

The new model is composed of four modules, each with a set of sub-models for: tree variable prediction, tree volume prediction, future tree list prediction and growth projection.

? Results

The evaluation of the PBRAVO and PBIRROL models showed that the new model gives more accurate predictions. Moreover, medium-term simulations provided consistent and logical predictions.

? Conclusion

It was verified that individual tree models are more suited to simulate poorly managed uneven-aged stands than diameter distribution models. No clear superiority of distance-dependent models was found over models using just distance-independent measures of inter-tree competition.     

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.

     

The effects of forest structure on the risk of wind damage at a landscape level in a boreal forest ecosystem

? The aim of this work was to analyze how the forest structure affects the risk of wind damage at the landscape level in a boreal forest.

? This was done by employing: (i) Monte Carlo simulation technique for generating landscapes with different age class distributions, proportions of open areas (gaps), and tree species composition; and (ii) a mechanistic wind damage model, HWIND, for predicting the critical wind speeds at downwind stand edges of open areas (gaps) for risk consideration. The level of risk of wind damage observed at the landscape level was significantly affected by the presence of gaps and old stands. Even a slight increase in the proportion of gap areas or older stands had a significant impact on the total length of edges at risk. As a comparison, variation in species composition (Scots pine and/or Norway spruce) had much smaller impact on the risk of damage.

? In conclusion, the effects of forest structure on the risk of wind damage should especially be considered by forest managers in day-to-day forest planning in order to reduce the risk of wind damage both at the stand and landscape level.

     

Effects of some ecological variables on carabid communities in native and non native forests in the Ibaizabal basin (Basque Country: Spain)

? Carabid beetles were investigated at five different forest types in the Ibaizabal basin (northern Spain). The landscape is characterized by the presence of remnants of native forest surrounded by conifer plantations.

? Carabids were trapped in 52 stands of mixed forest, beech forest, holm oak forest, mixed pine and Monterey pine plantations in 2005 and 2006. The main objectives of the study were: compare carabid diversity, recognise the characteristic species, and study the effects of ecological variables on carabid assemblages in the different forest types.

? No significative differences in species abundance, richness and diversity were found among the studied forests. Most of the trapped beetles were identified as forest generalists, nevertheless some native and non-native forest specialist species were also found. Distribution of carabid communities overlapped and, except for beech forest, no specific communities were distinguished. Altitude, percentage of grass coverage and temperature were the main variables influencing species distribution.

? The results suggest high habitat homogeneity, caused by regeneration in pine plantations of the indigenous understorey, and by poor habitat quality in native forest (patchy remnants enclosed in conifer plantations). This situation could explain the similar carabid diversity.

     

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.

     

Physical properties of wood in thinned Scots pines (Pinus sylvestris L.) from plantations in northern Spain

? The physical properties of wood and the associated variations within and between trees were evaluated by analysing 770 small specimens of clear wood from 11 Pinus sylvestris L. trees thinned from 3 plantations.

? Within-tree variations in basic density or volumetric shrinkage increased with cambial age and decreased with increasing ring width. The effect of the height in the stem on wood properties was considered indirect and height was not included as an explanatory variable in the mixed models proposed to estimate basic density and volumetric shrinkage.

? The models had random components for the intercept parameter and explained 52.5% of the total variance in basic density and 56% of the total variance in volumetric shrinkage. Linear shrinkage in the direction of the grain was extremely variable.

? Between-tree variation and between-plot variation in the physical properties of wood were high, considering that all trees sampled were growing in similar sites and stands. It would then be desirable to predict physical properties of wood on living trees in order to use the quality of wood as a criterion for timber tree selection in thinnings.

     

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.

     

Why and where do adult trees die in a young secondary temperate forest? The role of neighbourhood

? The density and identity of tree neighbourhood is a key factor to explain tree mortality in forests, especially during the stem exclusion phase.

? To understand this process, we built a logistic model for mortality in a spatially explicit context, including tree and neighbourhood predictors. Additionally, we used this model to build mortality risk frequency distributions. Finally, we tested this model against a random mortality model to predict the spatial pattern of the forest.

? Annual mortality rate was high for pedunculate oak (Quercus robur, 6.99%), moderate for birch (Betula celtiberica, 2.19%) and Pyrenean oak (Q. pyrenaica, 1.58%) and low for beech (Fagus sylvatica, 0.26%). Mortality risk models for pedunculate oak and birch included stem diameter, tree height, canopy position and neighbourhood. Mortality was affected by the specific nature of the neighbourhood showing a clear competitive hierarchy: beech > pedunculate oak > birch. Models based on random mortality and logistic regression model were able to predict the spatial pattern of survivors although logistic regression predictions were more accurate.

? Our study highlights how simple models such as the random mortality one may obscure much more complex spatial interactions.

     

Gender, neighboring competition and habitat effects on the stem growth in dioecious Fraxinus mandshurica trees in a northern temperate forest

? This study analyses the sex ratios, and the effect of gender, neighboring competition and habitat factors on the stem growth of dioecious F. mandshurica trees in the Changbai Mountains of North-Eastern China.

? The general sex ratio of the flowering trees did not significantly deviate from 1:1. The sex ratio varied among age classes. A female-biased sex ratio was found in the age classes of 65–75 years and 85–95 years, but departure from 1:1 was not significant in other age classes.

? An analysis of variance revealed that the growth rate was extremely sensitive to gender and tree size, as shown by a highly significant gender-related and size-dependent effect. Male trees had a faster stem growth rate than female trees, and the assumption is that trade-offs between reproductive and vegetative processes can induce the difference in stem growth rate between genders.

? Stem growth rates are positively affected by soil moisture and tree size for both genders at all considered neighborhood distances. The growth rates of female trees were negatively affected by neighboring competition from other female trees at all neighborhood distances between 1 and 10 m, but were not significantly affected by male trees. The growth rates of male trees were negatively affected by neighboring competition from male trees for neighborhood distances between 3 and 10 m, but were not significantly affected by female trees.

? These results suggest that intrasexual competition (male-male competition and female-female competition) may cause difference in stem growth for both genders. Intersexual differences in sensitivity to neighboring plants are considered to be an important factor driving gender-specific growth patterns.

     

Effects of stand density on ecosystem properties of subalpine forests in the southern Rocky Mountains, USA

? Mixed coniferous, subalpine forest communities in the Rocky Mountains are historically dense and have experienced infrequent, high-severity fire. However, many of these high-elevation stands are thinned for a number of perceived benefits.

? We explored the effects of forest stand density on ecosystem properties in subalpine forests in Colorado, USA, 17–18 y after forests were managed for timber.

? Forest structure significantly altered the composition and chemical signature of plant communities. Previously managed stands contained lower density of overstory trees and higher ground cover compared to paired reference stands. Foliar phenolic concentration of several species was negatively related to basal area of overstory trees. Furthermore, reductions in stand density increased total foliar phenolic:nitrogen ratios in some species, suggesting that gap formation may drive long-term changes in litter quality. Despite significant changes in forest structure, reductions in stand density did not leave a strong legacy in surface soil properties, likely due to the integrity of soil organic matter reserves.

? Changes in forest structure associated with past management has left a long-term impact on plant communities but has only subtly altered soil nutrient cycling, possibly due to trade offs between litter decomposability and microclimate associated with reductions in canopy cover.

     

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.

     

Epicormic sprout development in pruned coast redwood: pruning severity, genotype, and sprouting characteristics

? Young coast redwood (Sequoia sempervirens (D. Don.) Endl.) trees were pruned to various heights to examine the effect of pruning severity on epicormic sprouting. Seven separate stands were used with as many as six treatments per stand in coastal Humboldt County, California, USA.

? Epicormic sprout development was affected by pruning severity but primarily at the most severe pruning treatments that removed all but the branches in the top 15% of tree height. Less severe treatments produced sprouts but the number and size of these sprouts were comparable to unpruned trees.

? Natural clonal patterns were also used to explore patterns of sprouting between genotypes. Linear mixed-effects models were developed to predict sprouting frequency as a function of pruning severity while accounting for the nested data structure (i.e., stem sections sampled nested within genotypes within treatments within sites).

? Comparing variances attributed to each of these random effects indicated that at any level of pruning severity, differences in epicormic sprouting between genotypes and sites expressed soon after pruning had disappeared after six growing seasons. Epicormic branches were more common two years after pruning than six years indicating many branches were dying. Branches were more common in the middle of the pruned bole, possibly because of competition from basal sprouts and the expanding tree crown.

     

Species variables or environmental variables as indicators of forest biodiversity: a case study using saproxylic beetles in Maritime pine plantations

? To assess the sustainability of plantation forest management we compare two types of biodiversity indicators. We used the species richness of saproxylic beetles as a case study to test the “species” and “environmental” indicator approaches. We compared single species abundance or occurrence and deadwood volume or diversity as predictor variables.

? Beetles were sampled with flight interception traps in 40 Maritime pine plantation stands. The volume and diversity of deadwood was estimated with line intersect and plot sampling in the same stands. Predictive models of species richness were built with simple linear or Partial Least Square regressions.

? Deadwood variables appeared to be good predictors of saproxylic beetle richness at the stand-scale with at least 75% of variance explained. Deadwood diversity variables consistently provided better predictive models than volume variables. The best environmental indicator was the diversity of deadwood elements larger than 15 cm in diameter.

? By contrast, the use of “species variables” appeared to be less relevant. To reach the quality of prediction obtained with “environmental variables”, the abundance or occurrence of 6 to 7 species — some of which are difficult to identify — had to be used to build the indicator.

     

Density dependence on tree survival in an old-growth temperate forest in northeastern China

? Density dependence is a major mechanism for shaping plant communities. However, its role in regulating diverse, mixed natural tree communities is less certain.

? In this study we investigated density-dependent effects in a large-scale (25 ha) old-growth temperate forest in northeastern China. Spatial patterns of neighborhood distribution in the plot were analyzed using various methods for inferring competition, including (1) pair correlation function to determine spatial patterns of pre-mortality and post-mortality and (2) neighborhood analysis of individuals to examine the extent to which tree survival is correlated with other covariates.

? Results showed that, for common species, 3 of 5 canopy species and 3 of 8 midstory and understory species were random in mortality. Negative density-dependent mortality was not found when trees reach 1 cm in DBH. There was no significant correlation for canopy species between tree survival and conspecific abundance, but largely positive correlations for midstory and understory species. In contrast, tree survival was found to negatively correlate with conspecific basal area for most species, indicating strong intraspecific competition. No strong interspecific density dependence was found in the forest.

     

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.