Ranking larch genotypes with the Rigidimeter: relationships between modulus of elasticity of standing trees and of sawn timber

? Direct assessment of modulus of elasticity (MOE) on standing trees is attractive for breeders to evaluate genotypes prior to selection: this can be done using the Rigidimeter, a bending-based measurement device.

? In this study, we tested its reliability to properly rank genotypes by relating trunk MOE with MOEs estimated with a vibrating analysis system (Bing) on different types of conditioned wood specimens from the same trees (boards and standardised 2×2×30 cm-clear-wood specimens). One hundred and ten trees from different genotypes of hybrid larch (Larix × eurolepis) were tested.

? Mean trunk MOE was 7 300 MPa with a similar value obtained for sawn boards. Clear-wood specimens MOE increased from pith to bark from less than 6 000 MPa to nearly 9 000 MPa. Moderate correlations (r = 0.48–0.61) were found at the individual tree level between trunk MOE and MOE of wood samples.

? Single specimen MOE was shown to be strongly related to a linear combination of trunk MOE and sample position.

? At the genotype mean level, trunk MOE was highly correlated with wood samples MOE (r = 0.80–0.91). Ranking of genotypes based on trunk MOE was mostly consistent with that based on standardised specimens.

? It was concluded that besides other operational advantages which are discussed, the Rigidimeter is a valuable tool for breeders to routinely evaluate and rank genotypes for stiffness prior to further selection.

     

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.

     

Wood colour variation in sapwood and heartwood of young trees of Tectona grandis and its relationship with plantation characteristics, site, and decay resistance

? Wood colour of Tectona grandis produced from fast-growth plantations is highly variable and the causes of this variation are relatively unknown.

? With the purpose of understanding the colour variation, different fast-growth plantations were sampled with different growth rates, tree ages, and sites.

? Wood colour was measured with a CIELab system, where three variables are estimated: coordinate L* for lightness, coordinate a* defines redness and coordinate b* defines yellowness.

? Results showed only a negative correlation between L* and a*. L* and a* were negatively and positively respectively correlated with pith distance in heartwood, but not for b*. No correlations were found between L* and b* in sapwood and plantation characteristics, while a* was positively correlated with age and height of tree and growth rate. In heartwood, tree age and diameter at breast height were correlated with all colour parameters, but tree height and plantation density were correlated with a* and b*. Cluster site had correlation with L*. Multiple correlation analysis showed that the heartwood is increasing darker (L*) and redder (a*) when the trees are older and bigger. Correlation coefficient shown that sapwood and heartwood with lighter colour (L*) is less resistance to fungal attack, but redness colour (a*) increasing decay resistance.

     

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.

     

Stimulation of tree defenses by Ophiostomatoid fungi can explain attack success of bark beetles on conifers

? Our aim is to present why the hypothesis, that Ophiostomatoid fungi play an important role in the establishment of most bark beetle species on living conifers, is valuable.

? After summarizing knowledge about the relationships of bark beetles with conifers and fungi, we conclude that controversy results from misinterpretations when using fungal pathogenicity to demonstrate the role of Ophiostomatoid fungi in beetle establishment on host trees.

? We demonstrate that fungal pathogenicity is not the right parameter to appreciate the role of fungus in beetle establishment on host trees. We argue that artificial low density inoculations that allow the appreciation of fungus ability to stimulate tree defenses and thus to help beetles in overcoming tree resistance must be used in complement to mass inoculations. In both cases, results must be expressed in terms of tree defense stimulation rather than in terms of tree killing.

1. Fungal species stimulating tree defenses are generally not those that grow the best in the sapwood.
2. We argue that beetle development in the phloem, fungal invasion of the sapwood and phloem, and tree death, occur after tree defenses are exhausted, and that any fungus present in the beetle gallery could thus potentially invade the sapwood after defense exhaustion.

? We conclude that stimulation of the tree defense reactions in both the phloem and the superficial sapwood is a real benefit brought by fungi to the beetles during the first phase of establishment (overcoming tree resistance).

? Considering the origin of the bark beetle fungus associations attacking living trees and their general functioning based on stimulation of tree defenses, we develop three hypotheses:

1. any beetle species would be helped in its establishment in a given tree species by developing an association, even loosely, with a fungus species belonging to the Ophiostomatoid flora of that tree species;
2. the necessity of a considerably low level of tree resistance for fungus extension into the tree is the selection pressure that has led fungi to develop their intrinsic ability to stimulate tree defenses, through their ability to grow into the phloem. This association can be completed by antagonistic fungal species controlling extension of the previous fungal species in the tree tissues;
3. Beetle species using the strategy of overcoming tree resistance are associated with a fungal complex, of which species could assume three roles regarding relationships between beetles and trees: 1- to stimulate tree defenses in the phloem and superficial sapwood, 2- to grow into the sapwood after tree resistance is overcome, and 3- to control phloem extension of the first other two categories. Bringing nutrients to the beetle progeny can be a fourth role.

? We propose that bark beetle — Ophiostomatoid associations can be categorized, based on associations’ frequency and complexity while taking into account beetle aggressiveness. We show that a close correspondence exists between beetles’ aggressiveness and the ability of their main associated fungal species to stimulate the defenses of their host tree.

? We conclude with suggesting that most sapwood invading fungi might be “cheaters” which have taken advantage of the efficiency of the relationship between beetles and fungi that stimulate tree defenses.

     

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.

     

Extending sapwood — Leaf area relationships from stems to roots in Coast Douglas-fir

? Studies of allometric relationships between leaf area and the cross-sectional area (CSA) of sapwood in the stem have shed light on the structural and functional relationships between water-conducting and photosynthetic tissues.

? The purpose of this study was to test whether sapwood-leaf area relationships could be extended from stems to roots in coast Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirb.) Franco). Twelve trees were felled, their stumps were excavated, and the CSA of sapwood and heartwood were estimated for individual roots, entire root systems, and stem section.

? Root sapwood CSA was greater than sapwood CSA throughout the stem, and the ratio of leaf area to sapwood CSA (A _l :A _s ) was accordingly lower for root sapwood. The relationship between sapwood CSA and leaf area was more variable in roots and at groundline compared to crown base. Root A _l :A _s decreased with relative tree height (tree height/mean stand height).

? The strong allometric relationship between leaf area and the CSA of sapwood in the stem generally holds when extended to roots. The greater CSA of sapwood in roots versus stems may reflect differences in their roles in supporting the tree.

     

Colour and decay resistance and its relationships in Eperua grandiflora

? Eperua grandiflora, which is widely distributed in the French Guiana forest region, shows high variability in decay resistance. Further information concerning this wood quality parameter is necessary, but standard testing methods are complex and time-consuming. We assessed the use of colorimetry to determine durability in heartwood samples from a range of trees.

? Eperua grandiflora colour parameters were measured using a CIELAB system, revealing that the tree effect was greater than the radial position and height effects.

? The wood samples were exposed to Coriolus versicolor and Antrodia sp. according to two European standards (En 350-1 and XP CEN TS 15083-1). Eperua grandiflora is more susceptible to brown rot. These two standards did not give the same durability classes. The high variation in natural durability was due to the tree effect.

? These two properties were found to be correlated and the assessment also distinguished the extreme durability classes but they are not sufficient to classify the class of durability of this species.

     

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.

     

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.

     

Variation in the decay resistance of larch to fungi

? Decay resistance of larch (Larix sp.) to fungi was evaluated on heartwood samples belonging to 3 species (L. decidua, L. kaempferi and their hybrid), 3 races of European larch (polonica, sudetica and alpine), 13 wood lots (populations) and 313 trees.

? Larch wood appeared, on average, as moderately durable although a high variability was observed. At the sample level as well as at the mean individual tree level, durability ranged from class 1 to 5 according to EN 350-1 standard. At the population level, larch wood varied from ‘durable’ to ‘slightly durable’. Genetics played a major role in decay resistance at the species, provenance and tree levels. Environmental factors such as the position of heartwood samples and the age of trees were also identified as a source of variability.

? The most durable wood was not necessarily from old native alpine stands of European larch: some young larches from faster growing lowland origins also produced durable wood.

? Genetic improvement of larch wood durability appeared therefore likely by the selection of the best populations for decay resistance as well as from the selection of individuals.

     

Short-term effects of clear-cutting on the biomass and richness of epiphytic bryophytes in managed subtropical cloud forests

? Managed laurel forests in the Canary Islands have undergone clear-cutting with rotation periods of less than 30 y. Forest owners have recently requested a drastic reduction in the cutting interval. The effects of this new harvesting cycle on organisms like epiphytic bryophytes are not well known.

? This study investigates how time since last clear-cut, host species and characteristics of tree zones influence the biomass, cover and richness of epiphyte bryophytes in managed laurel forests in La Palma, Canary Islands. Four forest ages (8, 15, 25 and 60 y) and three host tree species (Erica arborea, Laurus novocanariensis and Myricafaya) were studied.

? Biomass, cover and richness of bryophytes increased through the chronosequence, both at the level of each plot and overall for L. novocanariensis. Most of the biomass (53%) and richness (81%) was concentrated in one of the tree species (L. novocanariensis), in plots for which 60 y had elapsed since the last clear-cutting. Trunks supported greater bryophyte biomass and richness than canopies, even in the oldest plots.

? Our results suggest that the current rotation periods used to manage laurel forests are insufficiently long to allow for complete reestablishment of epiphytic bryophyte assemblages.

     

Genetic parameters for early wood and latewood densities and development with increasing age in Scots pine

? Each annual ring in pines consists of earlywood and latewood with considerable difference in density and width. To get a better determination of the genetic regulation of total wood density in Scots pine (Pinus sylvestris L.), density and width of those ring sections were measured in annual ring numbers 12 to 21 of Scots pines in a full-sib progeny test. Tree height and stem diameter were also measured.

? Heritabilities for the annual ring sections increased with age for earlywood density from 0.08 to approximately 0.25; latewood density showed similar reductions. Heritability over all 10 annual rings was 0.25 for earlywood density, 0.22 for latewood density, 0.29 for height and 0.10 for stem diameter. Genetic correlations between earlywood and latewood density and growth traits were negative, while they were strongly positive between densities of adjacent annual rings (0.70–1.0).

? Despite the higher heritability of earlywood density, the strong positive genetic correlation between those traits indicates little benefit from focusing solely on earlywood density when selecting for wood density. Analysing earlywood and latewood separately does not benefit from including the width of the corresponding ring section as a covariate. Juvenile wood may possibly turn into mature wood 15–20 y from the pith.

     

Thigmomorphogenesis versus light in biomechanical growth strategies of saplings of two tropical rain forest tree species

? In the dense tropical rainforest understorey, saplings exhibit different growth strategies aiming at reaching light levels better fitting their ecology. Investing mainly in height growth, at the expense of their width, a lot are close to mechanical instability. Tachigali melinonii, a long living heliophilic tree species, is frequently observed to be extremely slender and supported by neighbours. Such observations suggest an active growth control through the perception of mechanical environment.

? Mechanical environment or light availability, which one is the most influent on growth and slenderness (H/D)? To test this question, we recorded growth of control and staked saplings of two species with contrasting habits and ecology: T. melinonii, and Dicorynia guianensis, along a natural light gradient.

? Dicorynia, the more stable, responded more clearly to the staking treatment, showing slenderness increase when light is available, whereas for Tachigali, only light availability governed growth.

? For Tachigali, growth allocation is mainly governed by light availability and ontogeny, whereas Dicorynia is probably similar to the average tree strategy, using the thigmomorphogenetic physiological process to control its stability.

     

Effects of climate variables on intra-annual stem radial increment in Pinus cembra (L.) along the alpine treeline ecotone

? Within the alpine treeline ecotone tree growth is increasingly restricted by extreme climate conditions. Although intra-annual stem growth recorded by dendrometers can be linked to climate, stem diameter increments in slow-growing subalpine trees are masked by changes in tree water status.

? We tested the hypothesis that intra-annual radial stem growth in Pinus cembra is influenced by different climate variables along the treeline ecotone in the Austrian Alps. Dendrometer traces were compared with dynamics of xylem cell development to date onset of cambial activity and radial stem growth in spring.

? Daily fluctuations in stem radius reflected changes in tree water status throughout the treeline ecotone. Extracted daily radial increments were significantly correlated with air temperature at the timberline and treeline only, where budburst, cambial activity and enlargement of first tracheids also occurred quite similarly. A close relationship was detected between radial increment and number of enlarging tracheids throughout the treeline ecotone.

? We conclude that (i) the relationship between climate and radial stem growth within the treeline ecotone is dependent on a close coupling to atmospheric climate conditions and (ii) initiation of cambial activity and radial growth in spring can be distinguished from stem re-hydration by histological analysis.

     

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

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

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

? Fecundity was high, with 138.2 eggs per female.

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

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

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

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

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

     

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.

     

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.

     

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.