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Open AccessArticle

Variation of Ring Width and Wood Density in Two Unmanaged Stands of the Mediterranean Oak Quercus faginea

1
Centro de Estudos Florestais, Instituto Superior de Agronomia, Universidade de Lisboa, Tapada da Ajuda, 1349-017 Lisboa, Portugal
2
Departamento Florestal/CITAB, Universidade de Trás-os-Montes e Alto Douro, Quinta de Prados Apartado 202, 5000-911 Vila Real, Portugal
*
Author to whom correspondence should be addressed.
Forests 2018, 9(1), 44; https://doi.org/10.3390/f9010044
Received: 27 November 2017 / Revised: 5 January 2018 / Accepted: 17 January 2018 / Published: 20 January 2018
(This article belongs to the Section Forest Ecophysiology and Biology)

Abstract

Ring width and wood density variation were studied from pith-to-bark and along the stem in two naturally regenerated stands of Quercus faginea Lam. in Portugal. Ring width was significantly different between sites, in both heartwood and sapwood rings, ranging from 1.83 mm to 2.52 mm and from 0.77 mm to 2.11 mm, respectively. Wood density was significantly different between sites only in the heartwood, i.e., 914 kg m−3 and 1037 kg m−3. Site effects were the main source of variation for ring width and wood density within the heartwood as well as for sapwood ring width, while the between-tree effects explained more the density variation within the sapwood. Wood density showed within-tree uniformity that was not affected by site. The stand characteristics such as basal area and tree age may override the environmental growth conditions. There was also a weak correlation between wood density and ring width components therefore suggesting the possibility of forestry management for both fast tree growth and high wood density.
Keywords: Quercus spp.; wood variability; wood characteristics; endemic species; sustainability Quercus spp.; wood variability; wood characteristics; endemic species; sustainability

1. Introduction

Quercus faginea Lam. is naturally distributed in the Iberian Peninsula and Maghreb Africa, in the Mediterranean Basin. It is commonly known as the Portuguese or Lusitanian oak and belongs to the white oaks subgroup. In Portugal, its present area is very restricted because of the past intensive utilization [1]. The species shows favorable characteristics for wood processing and performance, and is considered a good potential timber species for structural and flooring components, and cooperage [2,3,4,5].
Forest management practices combining wood production and conservation are gaining more interest and application in forestry and within ecological contexts, e.g., attention is given to mixed stand dynamics, slow growth species and low management conditions. In the case of Q. faginea, wood production doubled in mixed forests compared to mono-specific stands [6]. Regeneration of species, typically including slow growing and late-successional species such as deciduous oaks, suggested good performances for mixed-forests with fast-growing species such as Pinus spp. [7]. Knowledge of growth features over the tree‘s life-time, especially for slow growing species, as is the case for Q. faginea, will be important for better designing a forest management (e.g., stand location and density, rotation age and tree selection) that will include a high-value product-oriented approach.
The wood structure of Q. faginea is heterogeneous, as in other oaks with expected tree variability, namely of fiber and ray cells [5]. Despite the large number of relevant wood properties, the most used indicators of wood stem quality are ring width and wood density [8,9]. The typical radial ring width pattern in Q. faginea, and in oaks in general, is characterized by a decrease with cambial age (i.e., age from the pith) followed by some stabilization [10,11,12,13,14]. A decreasing trend in wood density is also common in Quercus spp. [10,11,15,16,17]. Wood density is related to both cambial age and ring width including ring structure, e.g., the latewood proportion is often the main responsible for wood density in ring-porous species [12,13,18]. Overall, the radial, pith-to-bark, variation is associated with the cambium maturation or physiological age [19,20].
It is widely known that oaks are highly appreciated due to their heartwood and wood technological characteristics related to high extractives content ensuring higher durability and attractive colors, as well as to specific features such as tyloses that provide less permeability and also increase durability [21,22,23]. Sapwood in general is undesirable for most timber uses and usually eliminated during the primary conversion. In Q. faginea, a positive relationship between heartwood proportion and tree diameter was found, i.e., 60–70% of heartwood for 20–25 cm stem diameters, while the sapwood width was relatively constant, ranging from 23 mm to 38 mm in older and younger trees, respectively [14].
The present study adds information on radial growth and wood density of Q. faginea based on trees from two different sites within the natural distribution range of this species. The specific aims of this study were to identify the main effects of site, trees, cambial age and stem height levels on wood variability of Q. faginea that could support a sustainable management including wood production.

2. Material and Methods

2.1. Sites and Sampling

The study was carried out in two locations, near Macedo de Cavaleiros (MC, northeast of Portugal), and near Vimeiro (VI, center of Portugal), both within the Q. faginea geographical natural distribution [1]. The stands resulted from natural regeneration, were unmanaged, mixed and uneven-aged, and VI is kept for state conservational purposes. The vegetation at MC is mainly characterized by minor occurrence of Q. suber L., Q. rotundifolia Lam. and Pinus pinaster Aiton trees, and mainly Cistus ladanifer L., Lavandula spp., Daphne gnidium L., Cytisus multiflorus (L’Hér) Sweet shrubs; at VI there were sparse Q. suber, Castanea sativa Mill. and P. pinaster trees, shrubs such as Arbutus unedo L., Ulex europeaus L., and Erica arborea L. and the fern Pteridium aquilinum L. (Kuhn).
The climate is of Mediterranean type at both sites. However, at Site MC the weather is described as interior Mediterranean characterized by dry and hot summer, and rainy winters (Csa) while at Site VI the coastal Mediterranean designation is applied with climate being characterized by cool and dry summer, and rainy winters (Csb) according to the international Köppen-Geiger climate classification system.
Basal area of Q. faginea was 18 m2/ha and 102 m2/ha, and overall stand density was 327 trees/ha and 300 trees/ha at MC and VI, respectively. Trees were on average 40 and 125 years old at MC and VI, respectively. Due to the reduction of Q. faginea distribution area it was allowed to cut only a very limited number of specimens. Ten dominant or co-dominant healthy trees were randomly selected from each site. Tree and site characteristics are shown in Table 1. From each of the 20 trees harvested, a disc was taken at stem base, 1.3 m, 3.4 m, and every 2.1 m along the stem to the top level.

2.2. X-Ray Microdensitometry

A radial sample from pith to bark was cut from each disc avoiding tension wood and knots. The radial samples were trimmed down to strips with 5 mm width and 2 mm thickness (longitudinal) with a specially designed dual-saw equipment, and then conditioned at 20 °C, 65% RH to 12% moisture content.
The radial strips were X-rayed perpendicularly to the transverse section with an accelerating tension of 12 kV, an intensity of 18 mA and a time of exposure to radiation of 350 s, at 2.5 m distance between X-ray source and Kodak film. The images were scanned by microdensitometric analysis and density was recorded at every 100 μm with a slit height of 455 μm. Detailed descriptions of the method are given in [24].
The ring boundary was identified on the transverse section by focusing on anatomical parameters, namely the pore distribution that is characterized by an abrupt size transition; and by visual cross-examination locating the sharp density variations set by the earlywood and latewood on the radial X-ray profiles.
Per each growth ring the following wood density and ring width components were obtained: ring density (RD), earlywood density (EWD), latewood density (LWD), the heterogeneity index (HI), ring width (RW) and latewood percentage (LWP). The within-ring determination of early and latewood density was made using the average of the minimum and maximum density values per each ring [3,25]. The heterogeneity index was calculated as the standard deviation of all density values across each ring to quantify the intra ring density variation [26].

2.3. Data Analysis

Analyses of variance were performed for wood density and ring width components to assess the effects of site (S), trees per site (T/S), cambial age or rings (R), stem height levels (L) and their interactions, according to the model presented in Table 2. The number of rings included in the analyses was limited by the uppermost height level of the youngest tree at Site MC. Based on previous research [3], two approaches were followed: (i) a core analysis of the trees made up with the innermost 15 rings with the same cambial age (vertical series, the heartwood); and (ii) a sheath analysis made with a sequence of the 10 outermost rings with the same chronological age (oblique series, in the sapwood).
The analyses were performed with three levels (base, 1.3 m, and 3.4 m of stem height) or four levels (including 5.5 m of stem height). Despite the differences regarding tree characteristics between sites the obtained results were qualitatively the same and therefore the four levels analyses were chosen to be presented here.
Correlation analysis was also performed between the studied variables.
The statistical analyses were performed using the JMP Statistical Software (SAS Institute Inc., Cary, NC, USA).

3. Results

3.1. Variation of Ring Width Components

Q. faginea growth rings were wider and latewood percentage was higher at Site MC with statistically significant between-site differences (Table 3).
Site effects were statistically more significant but explained less of the total variation of ring width in the core analysis (14%, P < 0.001) in opposition to the sheath analysis (49%, P < 0.01) (Table 4). The between-tree variation (T/S) was highly significant in both analysis (7–16%, P < 0.001) (Table 4). Overall, the ring width decreased from the base to the top (Table 5). However, the height level effects accounted little for the total variation of ring width even if its interaction with trees/sites (L × T/S) was highly significant (7–9%, P < 0.0001) (Table 4). In the initial period of tree growth, the effect of cambial age (given by the rings) and its interactions were mostly non-significant (Table 4). Within the sheath analysis, the effect of cambial age was highly significant but only accounted for 3% (P < 0.001) of the total variation (Table 4).
Latewood proportion variation was also mainly explained by site effects accounting for 11% (P < 0.001) and 12% (P < 0.001) of the total variation followed by trees/sites (T/S) accounting for 6% and 7% (P < 0.0001) within the core and sheath analysis, respectively (Table 4). Within the tree, the latewood proportion was quite constant axially at both sites (Table 5), e.g., the cambial age explained only 4% (P < 0.0001) of the total variation near the pith and the height level was non-significant in both analyses (Table 4).
Ring width and latewood proportion correlations were positive but only slightly significant at both sites (Table 6).

3.2. Variation of Wood Density Components

In the heartwood rings, the wood density was higher at Site VI but no differences were found in the sapwood rings (Table 3).
Site was the main source of variation for wood density components (RD, EWD and LWD) within the heartwood rings (30–35%, P < 0.001) but was non-significant within the sapwood rings (Table 4). The between-tree variability per site (T/S) was highly significant for both analyses accounting more for the total variation of wood density components in the sapwood (25–34%, P < 0.0001) than in the heartwood (16–21%, P < 0.0001). The interaction of height levels and trees/site (L × T/S) also showed similar results but explaining more the wood density variation in the sapwood rings (17–32%, P < 0.0001) than in the heartwood (12–14%, P < 0.0001) (Table 4).
The cambial age effects were non-significant in the sheath analysis; in the core analysis, cambial age was a highly significant effect (P < 0.001) but only accounted for 2–4% of the total variation (Table 4). There was a general axial decrease of wood density from the base to the top; in the core analysis, the differences were statistically significant although the height level effects only accounted for 3–5% of the total variation of the wood density components while in the sheath analysis they were non-significant (Table 4 and Table 5).
The wood produced at Site MC was more homogenous i.e., the heterogeneity index was lower than at Site VI, and site effects accounted for the major differences specially in the sapwood rings (35%, P < 0.001) (Table 3 and Table 4).
There were no negative correlations between wood density components and ring width or latewood proportion (Table 6).

4. Discussion

Environmental and stand conditions are main factors that account for ring width and wood density variability of Quercus spp. [9,12,13,19,27,28,29]. For example, in Q. petraea the water and nutrients soil availability and climate seem to have more effect in radial growth than in wood density [29,30].
In this study, however, as regards the soil conditions, higher growth rates would be expected at Site VI since Q. faginea prefers deeper soils with higher water availability [1,31], as is the case for Site VI (Table 1 and Table 3). More results were found on ring width that showed different correlations with temperature and precipitation suggesting that Q. faginea achieves a relative independence from the regional rainfall regime under aridity conditions [31,32,33]. Thus, other environmental characteristics present at Site MC may have been important to increase growth rates, e.g., low temperature and soil silica availability, that are related to good growth conditions for this species or even the evaporation rates. Moreover, the stand characteristics were obviously more favorable at Site MC where trees are younger and there is lower between-tree competition essentially due to a smaller basal area (Table 1). The present findings call attention to the link of increased radial growth with forest management as it was already reported for commercial oaks such as Q. petraea and Q. robur, by preferring shorter rotations and faster growth [12,29].
The present study also suggests that wood density variation is mainly explained by the site effects although interpretations must be done carefully due to small number of sites tested and stand age and density differences as pointed above. For example, in the more studied species such as Q. petraea, controversial results were found on the site effects evidence including no geographical and site quality effects at all [28,34]. In other species, e.g., Acacia melanoxylon, no between-site differences were found [35].
In general, higher wood density values are found in the heartwood in relation with the accumulation of heartwood extractives and eventual tension wood formation or presence of knots [9,36]. As tension and knot wood were avoided in the present measurements, it is the extractives content that should explain the density variations in Q. faginea wood between sites and between trees per site. However, the total heartwood extractives were approximately the same at both sites (19%) [4], suggesting that it does not explain per se the wood density variability found between these sites. Within-tree extractives content might be important to explain heart- and sapwood differences in wood density. In general, it was possible to detect a clear decrease in wood density at the heartwood/sapwood transition by visual observation of the studied samples and each wood density profile. Even if this tendency was more evident in older trees at Site VI and at the lower stem height levels, it emphasizes the extractives accumulation importance to explain wood density variability within tree. As suggested for Q. petraea [37], the high wood density in heartwood might be also related with changes in anatomical features during its maturation, namely the presence of rays and tyloses. Regarding sapwood, it was observed that its width was rather constant along the stem showing significant differences between sites [14], although this study showed that its wood density was not affected by site.
Tissues composition namely the latewood proportion affects ring width and wood density in Quercus spp. The constant latewood proportion trend found in Q. faginea was also observed in Q. petraea that is the axial variation following the same cambial rings, showed weaker relationships than following identical calendar rings [38]. Site and trees seemed to contribute more for its variation in the young Q. faginea trees development than in older trees. In Q. faginea, the positive correlation of latewood proportion with ring width was stronger than with wood density. These correlations were similar to those obtained for the European oak [3,12,18,27,39]. It might be suggested that latewood percentage seems to be less correlated to radial growth increase when other ecological or environmental effects are more important for its development in Q. faginea.
The correlations between ring width and wood density components were weak, as was already seen in previous studies [3,39]. Similar results were also found in 110-year-old Q. petraea trees at plot scale, although a correlation was present at tree scale, thereby suggesting that a better radial growth due to better site conditions does not imply higher wood density [29]. In other ring-porous species as Robinia pseudoacacia L. from different origins, the growth rate accounted for only a proportion of the variation in wood density and latewood proportion showed also inconsistent influence in wood density [40].
The explanation for the wood density differences between sites could be also related to other anatomical features, e.g., to the extent and size of the large earlywood vessels [5,37]. However, a recent study [41] showed similar earlywood vessel area and radial patterns of all vessel features at both sites, which suggests that other cellular features such as wood rays or latewood vessels might be involved.
Despite the site effects that might be also related with the different tree physiological stages, it is important to note that between-tree effects were the second main factor to explain wood density and ring width variability in Q. faginea, which is in agreement with other wood quality studies of the most commercial oaks [12,28,29].

5. Conclusions

In Q. faginea stands, the site effects seem to contribute more for wood density than for radial growth variability. Wood density was high regardless of site, ranging from 914 kg m−3 to 1037 kg m−3. Cambial age only accounted for 2–4% of the total variation of wood density components in the heartwood and it was not significant for ring width variability. The latewood proportion was constant within tree and not significantly correlated with wood density. The correlations between wood density and ring width, although positive at both sites, were weak.
This study suggests that Q. faginea stand conditions regarding basal area, density and tree age may override the site edaphic and climatic conditions stressing the importance of forest management for increased tree growth. However, further research on more sites with similar tree age is needed to clarify the effects of tree competition, environmental conditions, heartwood formation and anatomical features such as wood rays.
The results show the wood potential from the actual unmanaged stands of Q. faginea for high value wood components, thereby contributing to avoid the species decline through wood valorization and sustainable management.

Acknowledgements

CEF is a research unit funded by Fundação para a Ciência e a Tecnologia I.P. (FCT), Portugal (UID/AGR/00239/2013). The authors would like to thank S. Knapič for project management and S. Cardoso for logistic and fieldwork support and arrangements. Thanks are also due to A. Gonçalves for sample preparation and C. Fernandes for helping with density measurements. The first author acknowledges FCT for a fellowship (SFRH/BPD/97970/2013).

Author Contributions

V.B.S., J.L.L. and H.P. contributed to the general study conceptualization; J.L.L. and V.B.S. conducted and supervised the laboratory measurements and observations; J.L.L. performed the statistical analyses; and H.P. was responsible for funding acquisition and work supervision. All authors interpreted the data and contributed substantially to the manuscript preparation.

Conflicts of Interest

The authors declare no conflict of interest.

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Table 1. Description of the sampled Quercus faginea trees by each site. Mean of ten trees and standard deviation.
Table 1. Description of the sampled Quercus faginea trees by each site. Mean of ten trees and standard deviation.
Macedo de Cavaleiros (MC)Vimeiro (VI)
Latitude41°31′ N39°29′ N
Longitude06°51′ W09°01′ W
Altitude (m)540100
SoilOrthic Dystric and Eutric LeptosolsChromic Cambisols
Annual precipitation (mm)700 ± 141890 ± 249
Annual mean temperature (°C)12 ± 115 ± 3
Q. faginea basal area (m2/ha)18102
Stand density (trees/ha)327300
Tree height (m)10.5 ± 0.714.8 ± 2.3
Diameter (cm) *20.9 ± 4.236.7 ± 5.9
Crown height (m) **8.3 ±1.38.5 ± 2.3
Radius crown (m)2.4 ± 0.74.4 ± 1.1
Tree age ***40 ± 8125 ± 11
* Over bark diameter at 1.3 m of tree height; ** Crown height = Total tree height − branch-free stem height; *** Age based in ring counts at the stem base.
Table 2. Model used for analysis of variance for the ring width and density components.
Table 2. Model used for analysis of variance for the ring width and density components.
Source of VariationDegrees of FreedomExpected VarianceError Term
(1) Sites (S)s − 1σ2ε + rl σ2T/S + rlt σ2S(2)
(2) Trees/Sites (T/S)(t − 1)sσ2ε + rl σ2T/S(11)
(3) Levels (L)l − 1σ2ε + rts σ2L(11)
(4) L × S(l − 1)(s − 1)σ2ε + r σ2LT/S + rt σ2LS(5)
(5) L × T/S (l − 1)(t − 1)sσ2ε + r σ2LT/S(11)
(6) Rings (R)r − 1σ2ε + lts σ2R(11)
(7) R × S(r − 1)(s − 1)σ2ε + l σ2RT/S + lt σ2RS(8)
(8) R × T/S(r − 1)(t − 1)sσ2ε + l σ2RT/S(11)
(9) R × L(r − 1)(l − 1)σ2ε + ts σ2RL(11)
(10) R × L × S(r − 1)(l − 1)(s − 1)σ2ε + t σ2RLS(11)
(11) Residual (R × L × T/S)(r − 1)(l − 1)(t − 1)sσ2ε
T = trees (10/Site); L = height levels (4/tree); R = rings (15 or 10); σ2S, σ2T/S, σ2L, σ2LS, σ2LT/S, σ2R, σ2RS, σ2RT/S, σ2RL, σ2RLS and σ2ε are variance components due to Sites, Trees/Sites, Levels, Levels × Sites, Levels × Trees/Sites, Rings, Rings × Sites, Rings × Trees/Sites, Rings × Levels, Rings × Levels × Sites and Residual.
Table 3. Mean values (±standard deviation) of the ring width and wood density components measured within the core and sheath analyses at both sites. Mean of 10 trees per site and four stem height levels per tree (RD, ring density; EWD, earlywood density; LWD, latewood density; HI, heterogeneity index; RW, ring width; and LWP, latewood percentage).
Table 3. Mean values (±standard deviation) of the ring width and wood density components measured within the core and sheath analyses at both sites. Mean of 10 trees per site and four stem height levels per tree (RD, ring density; EWD, earlywood density; LWD, latewood density; HI, heterogeneity index; RW, ring width; and LWP, latewood percentage).
AnalysisSiteRD (g/cm3)EWD (g/cm3)LWD (g/cm3)HI (g/cm3)RW (mm)LWP (%)
CoreMC0.914 ± 0.114 a0.790 ± 0.148 a0.963 ± 0.103 a0.057 ± 0.039 a2.52 ± 1.27 a68.54 ± 14.11 a
VI1.037 ± 0.117 b0.965 ± 0.144 b1.076 ± 0.108 b0.085 ± 0.042 b1.83 ± 0.92 b60.25 ± 15.80 b
SheathMC0.751 ± 0.132 a0.611 ± 0.160 a0.827 ± 0.129 a0.114 ± 0.056 a2.11 ± 1.15 a63.01 ± 13.79 a
VI0.680 ± 0.131 a0.623 ± 0.148 a0.722 ± 0.130 b0.055 ± 0.050 b0.77 ± 0.47 b54.88 ± 15.83 b
Different letters for each variable and type of analysis (core, sheath) correspond to significant (P < 0.05) differences between sites by Duncan multiple test.
Table 4. Results of the core and sheath analysis of variance for the wood density and ring width components, showing their significance (P values) and the expected variation (EV%, percentage of total variation due to each source of variation), at both sites (RD, ring density; EWD, earlywood density; LWD, latewood density; HI, heterogeneity index; RW, ring width; and LWP, latewood percentage).
Table 4. Results of the core and sheath analysis of variance for the wood density and ring width components, showing their significance (P values) and the expected variation (EV%, percentage of total variation due to each source of variation), at both sites (RD, ring density; EWD, earlywood density; LWD, latewood density; HI, heterogeneity index; RW, ring width; and LWP, latewood percentage).
Analysis Source of VariationRD EWD LWD HI RW LWP
PEV%PEV%PEV%PEV%PEV%PEV%
CoreS0.000929.70.000234.50.00130.80.001713.80.000513.60.000811.4
T/S0.000119.60.000116.10.000120.80.000110.20.00017.10.00016.4
L0.00014.70.00014.40.00012.90.00014.80.00013.10.01210.6
L × S0.01564.20.02883.00.00934.80.15061.10.00156.20.28770.3
L × T/S0.000113.50.000111.80.000113.40.000110.60.00019.20.00015.9
R0.00012.30.00014.20.00011.90.00017.40.04240.50.00014.2
R × S0.00031.10.00011.60.05840.40.00013.40.01631.70.39070.1
R × T/S0.17740.50.15180.60.14370.60.16831.10.01513.20.51340
R × L0.00013.00.00012.20.00013.20.16480.50.00461.80.7830
R × L × S0.40060.10.36610.10.38650.10.09871.40.20171.00.7350
R × L × T/S 21.2 21.4 21.0 45.7 52.7 71.1
SheathS0.08348.10.75080.00.014219.70.000134.80.000149.30.001011.9
T/S0.000133.50.000124.50.000130.20.00018.00.000116.20.00016.5
L0.01990.40.06410.30.01040.40.00540.60.02120.10.14280.3
L × S0.20461.50.66220.00.08492.60.77680.00.75830.00.48740.0
L × T/S0.000123.60.000131.80.000117.10.000117.70.00017.20.00155.3
R0.12790.20.11930.30.54980.00.08070.30.00012.50.36600.1
R × S0.51090.00.51350.00.71150.00.26450.30.00972.10.10301.3
R × T/S0.23120.70.60630.00.17540.90.35280.40.000110.80.18212.2
R × L0.78590.00.97020.00.14620.40.48810.00.29010.10.52530.0
R × L × S0.15780.90.12731.40.24480.50.18980.90.46900.00.48590.0
R × L × T/S 31.2 41.7 28.2 37.0 11.6 72.5
Sites (S), Trees/Sites (T/S), Levels (L), Levels × Sites (L × S), Levels × Trees/Sites (L × T/S), Rings (R), Rings × Sites (R × S), Rings × Trees/Sites (R × T/S), Rings × Levels (R × L), Rings × Levels × Sites (R × L × S) and Residual (R × L × T/S).
Table 5. Mean values (±standard deviation) for the wood density components and ring width within the core and sheath analyses by tree height levels at both sites (RD, ring density; EWD, earlywood density; LWD, latewood density; HI, heterogeneity index; RW, ring width; and LWP, latewood percentage).
Table 5. Mean values (±standard deviation) for the wood density components and ring width within the core and sheath analyses by tree height levels at both sites (RD, ring density; EWD, earlywood density; LWD, latewood density; HI, heterogeneity index; RW, ring width; and LWP, latewood percentage).
AnalysisHeight (m)RD (g/cm3)EWD (g/cm3)LWD (g/cm3)HI (g/cm3)RW (mm)LWP (%)
Core5.5 0.943 ± 0.102 a0.842 ± 0.134 a0.998 ± 0.097 a0.080 ± 0.047 c1.9 ± 1.4 a62.1 ± 16.8 a
3.4 0.958 ± 0.124 b0.849 ± 0.165 a1.006 ± 0.116 a0.078 ± 0.046 c2.1 ± 1.1 b65.2 ± 14.6 b
1.3 0.980 ± 0.144 c0.881 ± 0.187 b1.022 ± 0.130 b0.070 ± 0.040 b2.3 ± 1.0 b65.3 ± 15.6 b
0.51.021 ± 0.137 d0.938 ± 0.177 c1.053 ± 0.125 c0.056 ± 0.034 a2.4 ± 1.0 c65.0 ± 14.7 b
Sheath5.5 0.717 ±0.120 ab0.618 ±0.146 ab0.782 ±0.120 bc0.089 ± 0.059 b1.41 ± 1.0 a58.2 ± 13.8 a
3.4 0.703 ± 0.134 a0.600 ± 0.135 a0.763 ± 0.147 a0.087 ± 0.062 b1.44 ± 1.1 b59.5 ± 14.7 a
1.3 0.728 ± 0.144 b0.624 ± 0.173 b0.786 ± 0.145 c0.085 ± 0.068 b1.39 ± 1.1 b60.5 ± 15.7 a
0.50.713 ±0.144 ab0.626 ± 0.160 b0.767 ±0.146 ab0.075 ± 0.055 a1.52 ± 1.1 c57.6 ± 17.1 a
Different letters for each variable and type of analysis (core, sheath) correspond to significant (P < 0.05) differences between height levels by Duncan multiple test.
Table 6. Correlation matrix (Pearson bivariate) for ring width and density components within the heartwood (lower triangle, n = 1200) and sapwood rings (upper triangle, n = 800) (RD, ring density; EWD, earlywood density; LWD, latewood density; RW, ring width; and LWP, latewood percentage).
Table 6. Correlation matrix (Pearson bivariate) for ring width and density components within the heartwood (lower triangle, n = 1200) and sapwood rings (upper triangle, n = 800) (RD, ring density; EWD, earlywood density; LWD, latewood density; RW, ring width; and LWP, latewood percentage).
Sapwood
Heartwood RDEWDLWDRWLWP
RD10.8930.9480.0610.099
EWD0.90410.772−0.128−0.163
LWD0.9690.84210.0870.027
RW−0.037−0.186−0.04410.385
LWP−0.013−0.287−0.0720.3711
Values statistically significant (P < 0.05) in bold.
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