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.

     

Spatiotemporal patterns in seedling emergence and early growth of two oak species direct-seeded on abandoned pastureland

? Fine-scale spatial and temporal establishment patterns of direct-seeded oaks on abandoned agricultural land have been little studied despite their potential importance for long-term stand structure.

? Here we periodically monitored seedling emergence and early growth of bur oak (Quercus macrocarpa Michx.) and red oak (Q. rubra L.) on an abandoned pasture, and tested the effects of herbaceous competition, rodents, and soil physicochemical properties.

? Herbaceous competition slightly decreased diameter growth, but rodents had little impact on establishment. Red oak seedlings emerged earlier than bur oak and in a greater proportion (92% vs. 56%). Seedling emergence and early growth of both species showed significant spatial structures that were partly explained by variation in soil physicochemical properties. Bur oak was more responsive to microenvironmental heterogeneity than red oak, yet much of the variation in emergence and growth of both species remained unexplained.

? This suggests that other factors, such as acorn size or genetic variability, may exert equal or greater control than microenvironmental heterogeneity over seedling emergence and early growth of these two oak species on abandoned pastureland.

     

Tree-ring Δ13C reveals the impact of past forest management on water-use efficiency in a Mediterranean oak coppice in Tuscany (Italy)

? The effects of thinning and heavy stand density reduction was investigated in Turkey oak (Quercus cerris L.) forests of central Italy, to evaluate the physiological responses and the growth status of trees that survived a past coppice cut and thinning to convert the stand to high-forest.

? The working hypothesis was that a strong decrease in stand density would cause a decreasing in canopy-intrinsic water-use efficiency (measured as the ratio of CO₂ assimilation to stomatal conductance, A/g), thus an increase in tree-ring carbon isotopic discrimination (Δ¹³C).

? The tree-ring Δ¹³C of the remaining trees (“survivors”) was found to have significantly (P < 0.05) raised between year two and year seven since the coppice stand was thinned (high-forest conversion thinnings). This effect was mostly caused by a large decrease in tree-rings Δ¹³C at control site which was characterized by high density and competition by trees. An increase in survivors tree-rings Δ¹³C probably indicates an improved water availability, possibly induced by a decrease in competition and in stand density or a decrease in the precipitation intercepted by the canopy (i.e., a stronger increase in g over A since a decrease in A is highly unlikely). A change in foliar nitrogen, foliar Δ¹³C and content in chlorophylls was also recorded seven years after thinning.

? Thinnings carried out to convert old abandoned coppices into high-forest stands induce short-term stimulation of Turkey oak growth by increasing light and water availability. We were able to make a detailed reconstruction of the impact of past silvicultural treatment on the stand using a tree-ring wood Δ¹³C time-series.

     

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.

     

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.

     

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.

     

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.

     

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).

     

Analysis of spatial patterns of oak decline in cork oak woodlands in Mediterranean conditions

? Cork oak mortality is a recurrent problem in southwestern Portugal. Despite the perception of increasingly visible damage in oak woodlands on drought-prone sites, the role of the various environmental factors in their decline is not clear.

? To describe the spatial patterns of cork oak (Quercus suber L.) mortality, a cork oak mortality index (MI) was determined for each landscape feature (agroforestry system, soil type, slope and aspect) using a GIS approach. To achieve this goal, a logistic regression model was formulated analyzing interactions between landscape attributes and allowing a prediction of cork oak mortality.

? Maximum values of MI were found in (i) shrublands and open woodlands with shrub encroachment (MI 6 and 3, respectively), where competition for soil water between tree and understory increases; and (ii) on lower slopes in the rounded hilltops and smooth hillsides or shallow soils where access to groundwater resources during summer drought is difficult.

? The model highlighted the importance of the agroforestry systems on cork oak mortality and may be used to identify sensitive areas where mitigation actions should be employed in a scenario of increasing drought severity in these Mediterranean ecosystems.

     

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.

     

Improving estimations of maximal stand density by combining Reineke’s size-density rule and the yield level, using the example of spruce (Picea abies (L.) Karst.) and European Beech (Fagus sylvatica L.)

? The Reineke Stand density rule relating stem numbers to the quadratic mean diameter is generally used as a reference for modelling maximal stand density.

? The linearity of this relationship after double logarithmic transformation is generally assumed, but it must be questioned for untouched stands and stands with a conventional thinning regime. Curvilinearity is demonstrated for some spruce and beech stands in Switzerland and shown to be statistically representative. This relationship is independent of the site index. It can be interpreted as a change in mortality in young stages mainly due to competition and in older stages more to ageing.

? A more accurate estimation of the maximal stand density needs to take into account the important variation around the mean course, known as the yield level. A simple method to assess the yield level of any stand regardless of whether it is thinned or not is presented, based on estimating the effect of a stand opening on the basal area.

     

Lifespan and mortality of old oaks — combining empirical and modelling approaches to support their management in Southern Sweden

? Old oaks (Quercus robur L.) play an important role in the southern Scandinavian landscape by providing habitat for a wide range of species, a large proportion of them being currently on the National Redlists.

? To provide support for the management of these trees, we review data on oak mortality and formulate a mortality-driven stochastic model analysing interactions between mortality rate, oak recruitment rate into 100–150 age class, and amount of oaks older than 200 years.

? Empirical annual mortality rates varied between 0 and 13% with average 1.68%. Trees older 200 years had an average mortality rate of 1.1%. Oaks in the high density forests showed higher mortality (3.2%) as compared to the trees growing in the low density forests (1.2%). A 400-year long modelling exercises indicated that under current mortality rates (regular mortality being centred around 1% annually; and irregular mortality 7% with average return time of 13 years) the long-term maintenance of 20 trees older than 200 years per ha would require an input rate of 1 to 5 trees × year^?1 × ha^?1 into the 100–150 years old class.

? The modelling highlighted the importance of initial oak abundance affecting amount of old trees at the end of shorter (100 years) simulation period.

     

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.

     

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.

     

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.

     

Phenotypic variation in the phenology of ascospore production between European populations of oak powdery mildew

? Oak powdery mildew severity (Erysiphe alphitoides) is usually mild in Europe because epidemics start late in spring, at the end of the first oak growth unit maturation. However, the disease can occasionally be very severe when strong infection occurs early during the development of the first growth unit, suggesting that host-pathogen synchrony in spring could be a critical factor in disease severity.

? We studied the timing of ascospore production in a given environment for four E. alphitoides populations sampled from SW France to the Czech Republic to determine whether this trait shows variation within Europe.

? Timing of ascospore production was clearly influenced by environmental factors as chasmothecia from a single origin showed very different dates of optimal ascospore production when transferred for overwintering in locations with different climate. In common garden experiments, no differences were observed between populations for the date of optimal ascospore production.

? Results suggest little genetic differentiation for timing of ascospore production for E. alphitoides populations across Europe and therefore a lack of local adaptation to their host phenology. Availability in ascospore inoculum is limited during host budburst, explaining the low infection usually observed on the first oak growth unit.

     

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.

     

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.

     

Potential use of pine plantations to restore native forests in a highly fragmented river basin

? In forests, the substitution of broadleaf species by conifers can reduce biodiversity because coniferous forests generally provide less diverse vascular understories than broadleaf forests. However, in some cases, former pine plantations might be useful for restoring native forests. We compared plant species composition on the plot scale in natural beech and mixed oak forests with that in plantations of Pinus radiata. Links between plant diversity and landscape parameters (patch size, fractal dimension and distance to the nearest patch of the same type) were investigated.

? The objective of this study was to evaluate the use of pine plantations for restoring native diversity in a zone where native forests are very fragmented.

? Similar to oak forests, plant diversity in pine plantations was high, mainly due to the presence of generalist species. Some species characteristic of oak forests also appeared in pine plantations, suggesting the onset of natural forest regeneration.

? These results suggest that pine plantations could be used to promote natural regeneration of original oak forests. Moreover, residual native stands should be conserved as important sources of native species and their seeds.