Oak powdery mildew changes growth patterns in its host tree: host tolerance response and potential manipulation of host physiology by the parasite

Context

Parasites can induce strong effects on their host’s growth, not only as a result of host resource exploitation (growth loss) but also with a potential adaptive value for host (tolerance response) and themselves (increased transmission).

Aims

We assessed these three types of phenotypic changes in oak seedlings infected by powdery mildew.

Methods

A manipulative field experiment with three levels of parasite inoculum was designed in order to tease apart infection from genetic effects on oak growth. Seedlings were monitored during 3 years for height growth, phenology and infection.

Results

Powdery mildew infection induced both significant growth loss and qualitative changes in plant architecture. The most striking and unexpected change was increased growth polycyclism in infected seedlings. This benefitted both the host as a form of compensation for infection-caused height loss, and the pathogen, by increasing sporulation.

Conclusion

The study highlights the effect of parasites in the expression of plant phenotypic traits, such as phenology and ultimately tree architecture. Both host tolerance and parasitic manipulation may be involved in the observed changes in growth patterns. These results suggest a complex interplay between development and defence in trees and emphasize the need to better assess tolerance mechanisms when considering the defence strategies of trees against pathogens.     

Tree species exhibit complex patterns of distribution in bottomland hardwood forests

?Context

Understanding tree interactions requires an insight into their spatial distribution.

?Aims

We looked for presence and extent of tree intraspecific spatial point pattern (random, aggregated, or overdispersed) and interspecific spatial point pattern (independent, aggregated, or segregated).

?Methods

We established twelve 0.64-ha plots in natural bottomland hardwood stands in the southeastern USA.

?Results

Spatial point pattern analyses (Ripley’s K, L, and L ₁₂) indicated that, when species were combined, trees were frequently aggregated and less commonly overdispersed. Plots with larger trees were more likely to exhibit overdispersion, confirming a shift to this pattern as trees grow. The intraspecific pattern of cherrybark oak and water oak was either aggregated or random. Sweetgum was aggregated on all plots and always at smaller distances (less than 5 m) than the two oak species. Intraspecific overdispersion was very rare. Interspecific segregation among the two oak species was more commonly observed (six plots) than aggregation (one plot). Cherrybark oak and sweetgum were segregated at some scale on seven of the 12 plots and aggregated on only two plots.

?Conclusion

The results from the analyses suggest that strong interspecific competition may result in segregation of trees from different species, while weaker intraspecific competition may lead to aggregations of conspecifics.     

Effects of pruning and stand density on the productionof new epicormic shoots in young stands of pedunculate oak (Quercus robur L.)

Context

Avoidance or control of epicormic shoots is among the major silvicultural challenges for the production of high-quality oak timber. In northern Europe, contemporary oak silviculture aims to produce valuable timber on a relatively short rotation, applying early, heavy thinning combined with artificial pruning.

Aims

The aim of this study was to analyse the effects of pruning and stand density on the production of new epicormic shoots on young trees of pedunculate oak (Quercus robur L.).

Methods

The study was based on two field experiments in even-aged stands of pedunculate oak subjected to different thinning practices and early selection of potential future crop trees. From ages 13 to 15 years, stem density was reduced to 300 trees ha^?1, 1,000 ha^?1 or stands remained unthinned. Pruning was conducted on selected trees at ages 22–24 years. At that age, the stem density in unthinned control plots ranged from 2,500 to 3,100 ha^?1. All treatments were replicated twice within each experiment.

Results

Pruning led to an overall increase in the total production of new epicormic shoots. More epicormic shoots were produced in the lower part of the stem (0–3 m in height) than in the upper part (3–6 m). The number of new epicormic shoots increased with increasing stand density.

Conclusion

Early, heavy thinning combined with high pruning at regular intervals may help shorten the rotation length for pedunculate oak without further reduction in wood quality than that which is caused by wider annual growth rings.     

Effect of selective logging on stand structure and tree species diversity in a subtropical evergreen broad-leaved forest

?Context

Selective logging followed by natural regeneration is rarely employed for restocking subtropical evergreen broad-leaved forests in East Asia compared with the use of clear-cutting.

?Aims

To clarify the succession of these forests, the effects of selective logging on stand structure, species diversity, and community similarity were studied in a mature and regenerating forest in Okinawa, Japan.

?Methods

Four study plots were established, and trees ≥1.2 m height were identified by species name, tree height, and diameter at breast height.

?Results

The results showed that the species composition of regenerating forest was similar to mature forest; however, the former had a greater species density and Shannon–Wiener index than the latter. Castanopsis sieboldii and Distylium racemosum, the predominant trees in the mature forest, continued to dominate the regenerating forest, with a broad layer distribution. High Sørensen and Jaccard community similarity indices for mature and regenerating forest indicated that the regeneration occurred in a progressive succession.

?Conclusion

The similar species composition and stand structure for both mature and regenerating forest, and the higher species diversity for the latter, provided no evidence of forest degeneration and suggested that the regenerating forest may develop into a stand similar to preselective logging forest.     

Intra- and interspecific competition differently influence growth and stem quality of young oaks (Quercus robur L. and Quercus petraea (Mattuschka) Liebl.)

Context

Cluster planting has become a conventional establishment method for oaks in Central Europe, where the spacing of seedlings within clusters varies between ‘nests’ (0.2?×?0.2 m) and ‘groups’ (1?×?1 m). Although the space between clusters is expected to fill with voluntary regeneration, its competitive effect on oak growth and quality had not been studied yet.

Aims

The aim of the study was to analyse the effects of inter- and intraspecific interactions on growth and quality of oaks grown in cluster plantings by quantifying the influence of neighbouring trees. In addition, we analysed whether the spatial position of oaks within groups (inner section or periphery) influenced their quality development.

Methods

Using Hegyi’s competition index, the influence of competition from intra- and interspecific trees from early, mid- and late-successional species, on diameter, height, slenderness and quality (length of branch-free bole) of 10- to 26-year-old oaks grown in cluster planting stands was quantified at seven sites in Baden-Württemberg and Hessen, Germany.

Results

In general, mid- and late-successional trees exerted a stronger competitive influence on growth of target oaks in clusters than the conspecific oaks and pioneer tree species. Oak quality development benefited from intraspecific competition, but self-pruning was not further promoted through additional interspecific competition. Within groups, inner oaks had a higher probability of developing into potential future crop trees than outer oaks.

Conclusion

Our study showed that intra- and interspecific competition had different effects on target oak trees and that these effect differed between nest and group plantings. The development of naturally regenerated and planted trainer trees in group plantings should be monitored carefully and if necessary be controlled through thinning or pollarding.     

Influence of tree size, reduced competition, and climate on the growth response of Pinus nigra Arn. salzmannii after fire

Context

After wildfire, surviving trees are of major ecological importance as they can help in the post-fire regeneration process. Although these trees may be damaged, they may also benefit from reduced fuel hazard and competition. However, little is known about the long-term growth response of surviving trees.

Aims

This study aims to explain short- to long-term variations in the postfire growth of surviving black pines in an area burnt in 1994, focusing on levels of fire severity and tree sizes.

Methods

Relative basal area increments were used to detect time-course variations in postfire radial tree growth depending on fire severity. Linear mixed-effects models were used to describe the factors affecting postfire ring growth.

Results

In the short term, fire caused stronger reduction in growth in small trees with increasing bole char height. However, as time since fire increased, a positive effect of fire on growth due to reduced competition counteracted the short-term fire impacts. Indeed, small surviving trees demonstrated a surge in growth 15 years after the fire.

Conclusion

It was concluded that reduced competition might offset the short-term negative effects of fire in surviving black pines.     

Summer drought and canopy opening increase the strength of the oak seedlings–shrub spatial association

? Context

The knowledge of how shrub–seedling interactions vary with summer drought, canopy opening, and tree species is crucial for adapting forest management to climate change.

? Aims

The aim of this study was to assess variation in shrub–oak recruitment associations along a south–north drought climate gradient and between two levels of canopy cover in coastal dune forest communities in a climate change-adapted forest management perspective.

? Material and methods

Mapped data of associational patterns of seedlings of three oak species with interspecific pooled shrubs were analyzed using a bivariate pair correlation function in 10 (0.315 ha) regeneration plots located in forest and recent gap sites along the climate gradient. An index of association strength was calculated in each plot and plotted against a summer moisture index.

? Results

The association strength increased with increasing summer drought from wet south to dry north and from closed forests to gaps.

? Conclusion

Consistent with facilitation theory, our results suggest that climate change may shift associational patterns in coastal dune forest communities towards more positive associations, in particular in canopy gaps. In a perspective of climate change, foresters may need to conserve understory shrubs in gaps in order to promote oak species regeneration.     

Snow gliding on a south-facing slope covered with larch trees

? Context

Snow gliding is a downhill motion of snow on the ground; observations have shown gliding to be possible not only on open slopes but also in forest stands. Larch stands, with their low canopy density and open forest structure with clearings and gaps, are particularly prone to high glide rates. Snow gliding may have negative effects on juvenescent trees which can be damaged by extraction from the ground.

? Aim

The goal of this study was to determine whether snow gliding depends on forest cover (canopy) and size of clearings.

? Methods

Snow gliding was measured during eight winter periods at six measuring positions (ranging from ‘dense forest’ to ‘open slope’) in and beside a larch stand in the Stubai Valley, Tyrol, Austria.

? Results

The results showed that gliding is strongly influenced by forest cover. Snow gliding increases with decreasing canopy density. The difference between the six measuring positions was highly significant (p?<?0.005).

? Conclusion

The identified glide cracks on at least two measuring positions, indicating extreme glide rates and, therefore, strong negative effects on juvenescent trees. To prevent glide rates of a magnitude such as this requires a mature forest with at least 300 stems/ha.     

Effect of nutrient removal on radial growth of Pinus sylvestris and Quercus petraea in Southern Germany

? Context

The rising demand of energy wood for heating purposes in Germany leads to concerns regarding the overexploitation of forests. A major aspect is the impact of whole-tree harvesting on long-term productivity of forest soils.

? Aims

This study aimed to analyze the effects of nutrient removal on productivity using the historically prevalent practice of litter raking. Since there is a lack of controlled whole-tree harvesting experiments in Germany, we used litter raking as a surrogate management practice entailing the removal of nutrients from forest stands.

? Methods

We used three sites with documented litter raking to analyze the effects of nutrient removal on productivity using dendroecological methods: two recent litter removal experiments in two Scots pine stands (Siegenburg and Burglengenfeld) and one oak stand (Eichhall) with documented historic litter raking. Basal area increment (BAI) and tree-ring characteristics were compared between periods with litter raking and the preceding periods for both treatment and control plots.

? Results

For the two Scots pine sites with a relatively short litter raking period, no effects of litter raking on BAI could be ascribed to nutrient removal. On the oak site with a longer history of litter utilization, the loss in BAI due to litter raking amounts to 22 % during the period with active raking and to still 17 % in the recovery period.

? Conclusions

These results contribute to the still very limited understanding about the impact of whole-tree harvesting on forest productivity in Germany by laying down an upper limit of possible effects due to nutrient removal, as nutrient loss by litter raking tends to be higher than nutrient loss by whole-tree harvesting.     

Within-population variability in architectural traits and suitability to forestry conditions in Nothofagus nervosa (= N. alpina; Nothofagaceae)

? Context

There is evidence that Nothofagus nervosa (= N. alpina) is suitable for timber production in temperate regions due to its wood quality and fast growth. However, high intra-specific variability in the architectural traits of this species limits its usefulness.

? Aims

This study was aimed at evaluating intra-specific variability in the architecture of N. nervosa trees at the population level, with emphasis on traits related to the suitability of trees for timber production.

? Methods

The size of 13-year-old trees installed in an experimental field population and the extent of differentiation between trunk and branches were compared among families (trees derived from different mother trees).

? Results

The large majority of trees exhibited architectural features indicative of high suitability for timber production: clear differentiation between trunk and main branches and few occurrences of permanent trunk forks, apex deaths, and sylleptic branching. Trees belonging to different families differed in height and trunk diameter. Micro-environmental conditions had a very significant effect on most architectural traits, including the trunk differentiation.

? Conclusions

N. nervosa emerges as a suitable option for timber production in temperate regions. Trunk growth seems to be under both environmental and genetic control in this species.     

Interspecific differences in the oleoresin production of Copaifera L. (Fabaceae) in the Amazon rainforest

? Context

Copaifera species produce an oleoresin of commercial importance that is widely extracted in Amazon communities.

? Aims

This paper addresses two questions. (1) What are the morphological characteristics of Copaifera species that influence oleoresin production? (2) How do different Copaifera species respond to repeated harvests?

? Methods

We performed a large-scale experiment in the Brazilian Amazon. We tapped 110 Copaifera trees belonging to four species, and several morphological tree characteristics were measured to determine their effect on oleoresin production.

? Results

The proportion of Copaifera reticulata and Copaifera paupera trees that yielded more than 1 ml oleoresin was higher than the other species. The oleoresin volumes from yielding Copaifera pubiflora trees were significantly higher than those from C. reticulata and C. paupera, with Copaifera multijuga yielding intermediate values. Interestingly, none of the studied morphological tree characteristics had a significant effect on the proportion of yielding trees. Hollowed trees yielded significantly smaller volumes than non-hollowed trees. Both the proportion of yielding trees and oleoresin volumes decreased from the first to the second harvests for C. reticulata and C. paupera; however, the opposite pattern was observed for C. pubiflora.

? Conclusions

Oleoresin production capacity varies by species, and management protocols should account for these differences.     

Modeling recent bark stripping by red deer (Cervus elaphus) in South Belgium coniferous stands

? Context

Over the past few decades, the impact of large herbivorous ungulates on forest vegetation has been clearly highlighted. Among those impacts, bark stripping of coniferous trees is one of the most damaging. Bark stripping leads to rot development, inducing serious loss of timber value.

? Aims

The present study aimed firstly at evidencing the factors explaining the variations observed in fresh bark peeling rate for spruce and Douglas-fir in southern Belgium and secondly at identifying the key factors to consider when setting up a deer management plan.

? Method

Fresh bark peeling rate was recorded with a systematic sampling survey from 2004 to 2007. The covered territory was then divided into 63 distinct hunting zones of area ranging from 1,000 to 25,000 ha. About 5,000 plots were monitored annually. Each zone was characterized with a large number of explanatory variables. The explanatory variables were integrated firstly into fixed linear models using a stepwise procedure, and then into a mixed model.

? Results

The significant variables included in the model (R ²?=?44 %) are (by decreasing order of importance) red deer densities, proportion of coniferous stands and agricultural areas, snow cover, distance to urban habitats, and species diversity in the understory.

? Conclusion

The models revealed the impacts of several factors on bark peeling: deer density, deer-carrying capacity of the territory, landscape structure, and severity of winter conditions. The adjusted model allowed subtracting the impact of winter conditions in order to produce a relevant indicator for hunting management. In addition, the model was used to assess the sensitivity of a forested area to bark peeling based on its environmental characteristics.     

Radial growth changes following hemlock woolly adelgid infestation of eastern hemlock

? Context

Hemlock woolly adelgid (Adelges tsugae) is an invasive insect that is defoliating and killing eastern hemlock (Tsuga canadensis) in the USA.

? Aims

We quantified changes in tree-ring growth rates and wood anatomy for living trees infested with hemlock woolly adelgid across six sites from Massachusetts (42°41′N) to Georgia (34°53′N) to identify growth responses of eastern hemlock that had survived infestation.

? Methods

Annual ring widths from infested eastern hemlocks were cross-dated and measured. Growth rates before and after infestation were compared. Two infested trees from Virginia were cut, and thin sections were prepared to identify changes in cell properties.

? Results

At three sites, trees experienced a significant decrease in radial growth after hemlock woolly adelgid arrival; however, the other three sites showed no change or increase in growth. Latewood produced after hemlock woolly adelgid infestation had significantly smaller cells with reduced cell wall thickness compared to latewood prior to infestation.

? Conclusion

At half the sites where hemlock woolly adelgid infested eastern hemlock trees were sampled, radial growth increased or remained unchanged. This unexpected response may be due to reduced competition due to mortality of other eastern hemlocks or physiological compensatory responses of increased photosynthetic rate and increased water use efficiency experienced by eastern hemlock infested with hemlock woolly adelgid.     

Thinning has a positive effect on growth dynamics and growth–climate relationships in Aleppo pine (Pinus halepensis) trees of different crown classes

? Context

Modification of stand density by thinning may buffer the response of tree growth and vigor to changes in climate by enhancing soil water availability.

? Aims

We tested the impact of thinning intensity on cambial growth of Aleppo pine (Pinus halepensis L.) under semi-arid, Mediterranean conditions.

? Methods

A multiple thinning experiment was established on an Aleppo pine plantation in Spain. We analysed the stem growth dynamics of two different crown classes under four different thinning intensities (15 %, 30 %, and 45 % removal of the basal area) for 2 years, based on biweekly band dendrometer recordings. Local relative extractable soil water was derived from the use of a water balance model Biljou© (available at https://appgeodb.nancy.inra.fr/biljou/) and used as an explanatory variable.

? Results

Radial growth was mainly controlled by soil water availability during the growing season, and differed by crown class. The growth rates of dominant trees were significantly higher than the growth rates of suppressed trees. Removal of 30 % and 45 % of the initial basal area produced a growth release in both dominant and suppressed trees that did not occur under less intense thinning treatments.

? Conclusions

Soil water availability was the main driver of radial growth during the growing season. Forest management confirmed its value for ameliorating the effects of water limitations on individual tree growth. These results may help managers understand how altering stand density will differentially affect diameter growth responses of Aleppo pine to short-term climatic fluctuations, promoting forests that are resilient to future climatic conditions.     

Coastal and interior Douglas-fir provenances differ in growth performance and response to drought episodes at adult age

Context

Since the 2003 drought and heat wave, Douglas-fir dieback has been reported in France in trees older than 30 years. Consequently, it is questioned whether selected Douglas-fir provenances are suited to the frequent and severe drought events which are forecast due to climate change.

Aims

Our objective was to contribute to the screening of variability in productivity and growth response to soil-water deficit of mature trees from provenances not currently used for plantation in France.

Methods

We sampled 22 provenances, including coastal and interior Douglas-fir, covering a wide part of its natural distribution, from Oregon to California for coastal provenances and from British Columbia to New Mexico for interior provenances. These provenances were planted at the mid 1970s in two provenance trials located in the south-west area of France. Variability of productivity, of wood density, and of radial growth in response to drought episodes among provenances was quantified and related to soil-water deficit computed by daily water balance calculations.

Results

Whatever the provenance, annual radial growth is highly dependent on local soil-water deficit (Felines R ² = 0.57, Le Treps R² = 0.49). Radial growth and wood properties exhibit large differences between provenances at 30 years old. Variability between provenances for all wood characteristics studied is mainly structured geographically. Coastal provenances perform best for productivity at 30 years old (619 cm²?±?59), and exhibit a small growth reduction in 2004, the second successive year of drought (?10.7 %?±?3.8). Surprisingly, the southern interior provenances from the driest environments in the natural range show a large growth reduction in 2004 (?30.5 %?±?5.2).

Conclusions

The provenances tested exhibited significant differences in growth performance and growth reduction induced by the soil-water deficit. The approach coupling retrospective analysis of radial growth on mature trees and water balance modelling is relevant for evaluating provenance adaptation to more frequent or severe drought episodes.     

Effects of region, soil, land use, and terrain type on fuelwood properties of five tree/shrub species in the Sahelian and Sudanian ecozones of Mali

? Context

There is little information about inter- and intra-specific variation in fuelwood properties of trees/shrubs in West Africa.

? Aims

We studied variation in fuelwood properties of Balanites aegyptiaca, Combretum glutinosum, Guiera senegalensis, Piliostigma reticulatum, and Zizyphus mauritiana in the Sahelian and Sudanian ecozones of Mali.

? Methods

Trees were sampled on different soil (sandy, sandy/loam, or rocky), land use (parkland agroforest or woodland), and terrain (flat, temporarily flooded, or hill slope) types in five regions extending from the drier eastern to the more humid western parts of Mali. Basic density, volatile matter, fixed carbon, ash content, moisture content, gross calorific value, and fuel value index were measured for each tree and adjusted for tree age.

? Results

All fuelwood properties differed significantly among species, but there were significant species by region interactions. Most fuelwood properties of four species differed significantly among regions. Soil and/or land use type had significant effects on a few fuelwood properties of four species.

? Conclusion

In general, fuelwood production is recommended in all regions for G. senegalensis, in drier regions for B. aegyptiaca and C. glutinosum, in more humid regions for Z. mauritiana, and on rocky soils for all species. Fuelwood production of P. reticulatum is not recommended.     

Genotypic parameters of wood density and fiber traits in triploid hybrid clones of Populus tomentosa at five clonal trials

? Context

A clear understanding of the genetic control of wood properties is a prerequisite for breeding for higher wood quality in Populus tomentosa Carr. hybrid clones.

? Aims

The experiments aimed at unraveling genetic and environmental effects on wood properties among triploid hybrid clones of P. tomentosa.

? Methods

We used 5-year-old clonal trials established in Northern China to assess the heritability of wood density and fiber traits. Two hundred seventy trees from nine clones were sampled in five sites.

? Results

Site had a very significant effect on all recorded traits. Despite this large site effect, a tight genetic control was detected and clonal repeatability varied between 0.53 and 0.95. Significant genotype?×?environment interactions were detected for most of the traits. Moderate to tight correlation were evidenced among traits but they were not consistent with that in several cases that were site-dependent.

? Conclusions

Our results revealed a tight genetic control over several wood properties and therefore breeding programs might be able to improve wood density, fiber length, and coarseness in these hybrids.     

Spatially explicit competition in a mixed planting of Araucaria cunninghamii and Flindersia brayleyana

? Context

A 20-year-old Nelder wheel planted with hoop pine (Araucaria cunninghamii Aiton ex D.Don) and Queensland maple (Flindersia brayleyana F.Muell.) in 18 spokes and 8 rings represents nominal point densities of 3,580, 2,150, 1,140, 595, 305, 158, 82, and 42 stems/ha and offers an opportunity to examine competition and spatial interaction between these two species.

? Aims

This study aimed to evaluate the intraspecific and interspecific competition between two contrasting tree species and to determine the distance over which competition can be observed.

? Methods

Competition was estimated using Hegyi’s index, implemented using the Simile visual modeling environment, and calibrated using nonlinear least squares with PEST.

? Results

Interactions were detected between pairs of stems closer than D _ij ?<?40(d _i ?+?d _j ) where D is distance (in centimeters) and d is stem diameter (in centimeters diameter at breast height). F. brayleyana trees surrounded by A. cunninghamii trees experience negligible competition, whereas A. cunninghamii surrounded by F. brayleyana trees suffer strong competition.

? Conclusion

Forty times diameter offers a useful guide to the extent of competition in even-aged stands planted with these species. Competition can be observed empirically when pairs of trees are closer than 40 times the sum of their diameters, but the intensity of the competition may vary considerably with species.     

Variation of wood color parameters of Tectona grandis and its relationship with physical environmental factors

? Context

Teak??s wood color is considered an important attribute in the marketing phase and it has been influenced by environmental setting, stand conditions and management, plant genetic source, and age. However, there is a lack of understanding about how the environmental factors might affect the teak??s wood color planted in short-rotation forest plantations.

? Aims

The aim of this study is to understand the relationship, gathered from generated information, between edaphic and climatic variables and their effects in the wood color variation of Tectona grandis from trees in forest plantations.

? Methods

Twenty-two plots were grouped in five cluster sites that shared similar climatic and soil conditions. Data about soil??s physical?Cchemical properties and climatic variables were collected and analyzed. Representative trees were harvested next to each plot in order to obtain a wood sample per tree at a diameter breast height. Wood color was measured using standardized CIELab??s chromaticity system.

? Results

After comparing the wood change color index (?E*) in the five studied clusters, it was found that heartwood produced from drier and fertile sites had more yellowish-brown color. The heartwood b* color index resulted with significant correlations (R?>?0.5, P?<?0.05) among nine climatic and eight edaphic variables.

? Conclusion

It was concluded that climatic variables should be considered as the first-order causal variables to explain wood color variation. Hence, darker b* wood color was associated with dry climates; also, with deeper and fertile sites.     

Variations of wood basic density with tree age and social classes in the axial direction within Pinus massoniana stems in Southern China

Context

Reliable estimates of wood density (WD) within individual trees could maximize the value of Pinus massoniana for specific end-use.

Aim

We examined and quantified the axial patterns of WD in trees with different social status in the stands.

Methods

Wood disks were sampled at the bottom, breast height, and middle of each 1-m sections from 108 stems, harvesting from three social classes in subtropical forests. A mixed-effects model was designed to quantify axial variation.

Results

The WD at different height was significantly different from the whole-stem WD (WWD) except the relative height of 0.1. An overall decrease of 133.8 kg m^?3 in WD was found from stem base to top. WD was significantly influenced by relative heights, tree age, and social class. WD of each relative height in mature trees was significantly higher than that of younger trees. Tree social class can affect WD development in the axial direction at age classes 2 and 3. Combining the fixed plus random effects, the final model explained 91 % of the observed variation in WD.

Conclusion

The WD development patterns in the axial direction vary considerably among tree age, diameter at breast height, and social class. To distinguish and supply timber for specific end-uses, we should use the axial variation in disk WD (DWD) instead of WWD directly. The accurate predictions of WD provided by the model could be used to optimally classify logs into different product classes and maximize economic benefits. We can use DWD at the relative height of 0.1 instead of WWD of a single tree.