34
Draft Integrated System of Equations for Estimating Stem Volume, Density and Biomass for Australian Red Cedar Plantation Journal: Canadian Journal of Forest Research Manuscript ID cjfr-2016-0135.R3 Manuscript Type: Article Date Submitted by the Author: 20-Jan-2017 Complete List of Authors: Calegario, Natalino; Federal University of Lavras, Forestry Gregoire, Timothy; Yale University Silva, Tatiane; Universidade Federal de Lavras, Forestry Tomazello Filho, Mario; University of Sao Paulo, Department of Forest Sciences Alves, Joyce; Universidade Federal de Lavras Keyword: Wood density, </i>Toona ciliata</i>, System of equations, Wood Technology, biomass estimation https://mc06.manuscriptcentral.com/cjfr-pubs Canadian Journal of Forest Research

Integrated System of Equations for Estimating Stem …...Draft 1 1 Integrated system of equations for estimating stem volume, density and biomass 2 for Australian red cedar plantations

  • Upload
    others

  • View
    3

  • Download
    0

Embed Size (px)

Citation preview

Draft

Integrated System of Equations for Estimating Stem

Volume, Density and Biomass for Australian Red Cedar Plantation

Journal: Canadian Journal of Forest Research

Manuscript ID cjfr-2016-0135.R3

Manuscript Type: Article

Date Submitted by the Author: 20-Jan-2017

Complete List of Authors: Calegario, Natalino; Federal University of Lavras, Forestry

Gregoire, Timothy; Yale University Silva, Tatiane; Universidade Federal de Lavras, Forestry Tomazello Filho, Mario; University of Sao Paulo, Department of Forest Sciences Alves, Joyce; Universidade Federal de Lavras

Keyword: Wood density, </i>Toona ciliata</i>, System of equations, Wood Technology, biomass estimation

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

1

Integrated system of equations for estimating stem volume, density and biomass 1

for Australian red cedar plantations 2

Natalino Calegario, School of Forestry and Environmental Studies, Yale University, 3

New Haven, CT 06511, USA. (e-mail: [email protected]) 4

Timothy G. Gregoire, School of Forestry and Environmental Studies, Yale University, 5

New Haven, CT 06511, USA. (e-mail: [email protected]) 6

Tatiane Antunes da Silva, Department of Forest Science, Federal University of Lavras, 7

Lavras, MG 37200, Brazil. (e-mail: [email protected]) 8

Mario Tomazello Filho, Department of Forest Sciences, College of Agriculture “Luiz de 9

Queiroz”, University of São Paulo, Piracicaba, 13418, Sao Paulo, Brazil (e-mail: 10

[email protected]) 11

Joyce A. Alves. Department of Forest Science, Federal University of Lavras, Lavras, 12

MG 37200, Brazil. (e-mail: [email protected]) 13

Corresponding author. Name: Natalino Calegario; e-mail: [email protected]; Phone: 1 14

203-219-1052; Mail: 195 Prospect Street, Yale University, New Haven, CT, USA 15

06511. 16

17

18

19

20

21

22

23

24

25

Page 1 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

2

Abstract 26

A system of equations is proposed to assess the stem wood density variation of Toona 27

ciliata M. Roem. stems growing in Brazilian plantations. As a taper function, a 3-degree 28

polynomial was fitted and the stem radius squared (r2), the dependent variable, was 29

estimated as a function of diameter at breast height (dbh), total height (ht) and radius (r) 30

at height (h). A nonlinear function was fitted to estimate wood density variation, having 31

as the independent variable the ratio between radius r and height h. The stem mass was 32

estimated by integrating the product of stem volume and wood density. Stem 33

measurements from a group of 72 trees of Toona ciliata M. Roem. were used to fit the 34

taper equation. A group of 6 trees was selected and, using x-ray technology, a wood 35

density database was created. Both the taper and the nonlinear functions performed well 36

in estimating the radius and the wood density. The within tree wood density 37

systematically increased from pith to bark and from the base to the top of the tree. With 38

the density varying from base to top, the estimated mass of the stem, compared to the 39

mass estimated using wood density value at dbh, had bias of 4.2%. When the density 40

variation from base to top and from the pith to the bark of the tree was considered, the 41

estimated mass had a bias of 1.5%. 42

Key words: wood density, Toona ciliata, system of equation, wood technology, biomass 43

estimation. 44

45

Introduction 46

As the stem mass can be estimated by the product of stem volume and wood 47

stem density, the individual stem and stand mass estimation strongly depends on the 48

individual stem and stand volume and wood density estimations. More precise stem 49

volume and density estimation provides more precise mass estimation. 50

Page 2 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

3

Toona ciliata is the most wide-ranging of the four Toona species, occurring 51

naturally across much of South and Southeast Asia – from Pakistan and western China 52

to Indonesia and Malaysia – and across parts of Oceania, including Australia and 53

islands in the western Pacific Ocean (Hua and Edmonds 2008). Its timber is used for 54

high quality furniture, carvings, decorative panels, veneers, flooring, tea-chests, oil 55

casks, musical instruments, other decorative objects (Edmonds 1993). Also, it was 56

especially prized for carriage building in Australia and has been widely used in many 57

parts of the world for general house, vehicle, canoe, oar and boat construction (Vader, 58

1987, cited by Edmonds, 1993). 59

Individual stem volume and taper has been focus of a wide range of studies. The 60

first registered publications in form class volume and taper tables are credited to Höjer 61

(1903), Schiffel (1905) and Jonson (1910), cited by Behre (1927). Schumacher and Hall 62

(1933) presented a mathematical expression relating total and merchantable stem 63

volumes with dbh and total height. Many studies have been published subsequently in 64

which stem taper is modeled, which in turn leads to the estimation of stem volume, 65

following the evolution in statistic/mathematical methods and computation. 66

Publications such as Furnival (1961), proposing an index to compare equations for 67

individual tree volume estimation with different dependent variable dimension, Kozak 68

et al. (1969), using two-degrees polynomial taper equation in inventory estimating and 69

Demaerschalk (1973), proposing an integrated system for Tree Taper and volume 70

estimation, are considered pioneers in taper and individual tree modeling. Using a more 71

sophisticated approach, Gregoire and Schabenberger (1996) developed a nonlinear 72

mixed-effects model to predict bole volume of yellow poplars standing trees. In general, 73

the individual tree volume function is capable of estimating either total or partial stem 74

volume, inside and outside bark. The partial volume estimation is more flexible and 75

Page 3 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

4

allows multiproduct estimations, such as saw wood, pulp wood, and fuel. The 76

multiproduct estimations are performed by fitting a taper function and, using solid of 77

revolution techniques, the taper function is integrated within a height interval to obtain 78

the partial stem volume. In situations where the average wood density is available for 79

the interval the interval’s mass could be estimated unbiasedly. Furthermore, the 80

availability of a mathematical relationship between mass and carbon also will generate 81

model-unbiased carbon estimation. 82

By definition, the stand timber mass depends on the stem volume and density. 83

The stem wood density is the single most important wood property because its strong 84

relationship to both yield and quality as well as its large variance and heritability 85

(Zobel, 1963). Within the tree, wood density varies from base to top and from pith to 86

bark. Wood density varies because of changes in vessel size and frequency, fiber 87

dimensions (wall thickness and diameter), the percentage of parenchyma (e.g. ray) and 88

wood chemistry (Downes et al. 1997). Luxford & Markwardt (1932), studying the 89

strength and related properties of redwood, found that the specific gravity in redwood 90

stems varied laterally and vertically, and greater values were related to greater crushing 91

strength, side hardness and modulus of rupture. 92

The wood density variation vertically has motivated of study for many other 93

researchers . Zobel and van Buijtenen (1989) cited Chidester et al. (1938) as the pioneer 94

publication in wood density variation. In the study carried out in Eucalyptus regnans in 95

New Zealand, Frederick at al. (1982) found a decline in wood density from the tree base 96

to 1.4 m, with the lowest values occurring at 1.4 m, and higher than that the wood 97

density increases. 98

Also, from pith to bark density variation has been investigated in many research 99

studies. Heitz et al. (2013), collecting wood samples from two neotropical rain forests, 100

Page 4 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

5

found that the wood density tends to increase towards the outside in trees with low 101

initial density and to decrease towards the outside in trees with high initial density. For 102

Pinus radiata, the major source of variability in wood density at dbh was between trees 103

within each site (Raymond and Joe 2007). In Norway Spruce stems, the largest (49–104

80%) variation in wood density was found within the annual rings (Jyske et al. 2008). In 105

Ochroma pyramidal, the specific gravity increased linearly with radial distance at any 106

given height (Rueda and Williamson, 1992). In Eucalyptus globulus trees, sampled at 107

different heights, the wood density increased from base to top of the tree (Quilhó and 108

Pereira 2001). In Chestnut, when moving from the first ring width class to the seventh 109

class, a total decrease in specific gravity of 12.7% was observed (Romagnoli et al., 110

2014). 111

Modeling stem taper and tree specific gravity, Parresol and Thomas (1989) 112

applied the density-integral approach to estimate stem biomass for slash pine and 113

willow oak, considering the specific gravity as a function of height on the stem and age. 114

Using the Parresol and Thomas model, Gregoire et al. (2000) studied the statistical 115

properties of volume prediction from an integrated taper equation and derived the first 116

two moments of the volume predictor and the prediction error. 117

Even though many studies have focused on taper models and their integrals, 118

wood density, and tree and stand mass, nobody has yet developed an unbiased 119

integrated equation system to estimate stem mass by including base to top and pith to 120

bark wood density variation. So, the main objective of this research is to develop a 121

flexible system of equations to generate unbiased estimates for total or partial tree stem 122

volume, density and mass. In addition, the system is capable of evaluating the wood 123

density variation from the base to the top of the tree and from pith to bark. 124

125

Page 5 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

6

126

Materials and methods 127

128

Study area and trees sampled 129

The study area is located in the Campo Belo County, Minas Gerais state, Brazil. 130

The Köppen climate classification is Cwa, temperate humid, with dry winter and rainy 131

summer (Köppen, 1900). The warmest month exceeds 22 ° C, with altitude ranging 132

from 790 m to 1146 m and average rainfall of 1250 mm (INMET, 2016). A progeny test 133

was established on February 6, 2008, containing 78 progenies of half-brothers, which 134

were formed from seeds collected in open pollinated arrays in Queensland and New 135

South Wales, Australia. In order to test the progeny performance, a complete block 136

experimental design was installed with 3 replications and 16 trees per sampling unit, 137

and the tree spacing of 3 x 3 m. 138

Selection of trees 139

Two progenies, from Queensland, with better performance were selected and a 140

group of 72 trees were felled (34 trees for the first progeny and 38 for the second 141

progeny), with a mean age of 4.7 years old, based on the normality shape of the 142

diameter (dbh) distribution, i.e 68% of the trees between -1 and 1 standard deviation 143

around of the mean, 95% between -2 and 2 standard deviations and 99% between -3 and 144

3 standard deviations. Using a systematic subsample design, the upper stem diameters, 145

inside and outside bark, were measured at the following heights: 0.15 m (base), 0.7 m, 146

1.3 m (dbh), 30%, 50%, 70% and 85% of the total height. This data were used to fit the 147

taper function. Also, based on the diameter distribution of the felled trees, 6, of the 72 148

trees, were selected for measurement by X-ray densitometry, which generated the 149

Page 6 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

7

apparent wood density data base. The small size of the subsample of tree was offset by 150

the extensive series of measurement that were taken with each stem. approximately 151

10,000. This was a laborious and expensive process. . The 6 trees were felled and the 152

total height (ht) was measured, using a tape, and samples of 4 cm-disks were taken from 153

the same position as the upper stem diameter measurements, generating a total of 7 154

disks for each tree. Each disk was identified by block, progeny, and tree number. The 155

disks were placed in a fresh air site for drying and subsequently cut and analyzed. 156

Wood density measurements 157

The wood samples were extracted from the sample disks, and density was 158

assessed from pith to bark; each sample was 4 cm wide, 1.6 mm thick, across the radius 159

of the disk. The samples were stored in acclimatized chamber (20 °C, 60% RH) until 160

reaching the moisture content of 12%. The samples were placed on Tree Ring Scanner 161

equipment (QTRS-01X, Quintek Measurement Systems). Knowing moisture content is 162

important since the water present in the chemical components is critical to determining 163

the wood density (QMS, 1999). Since the sample humidity was kept at 12%, the wood 164

density measurements are called Apparent Wood Density (AWD). The equipment 165

scanned the samples and recorded the wood density values at intervals of 0.004 cm 166

apart, from pith to bark. 167

168

Geostatistical Analysis 169

Kriging was used to visualize density variation in a radial and longitudinal 170

approach. This technique, proposed by Krig (1951) and formalized by Matheron (1962), 171

consists in estimating the value z(k) among observations in different spatial locations. 172

Page 7 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

8

The observations that are close are more correlated than those are far away, quantified 173

as spatial autocorrelation. In a general geostatistic approach, the following variables are 174

defined: u=vector of spatial coordinates (in our case are the stem positions where the 175

apparent density values were observed); z(u)=apparent density values observed at u 176

position; k = position to be estimated z(k) value; h=lag, or spatial distance, between k 177

and each u spatial location; z(u+h)=lagged version of apparent density; N(h)=number 178

of points separated by lag h. z(u) is called tail variable and z(u+h) is referred to as the 179

head. After the lag vector is determined, the covariance, correlation and semivariance 180

are estimated and the parameters range, sill and nugget are calculated from 181

semivariance function. Based on semivariance parameters, a semivariance model is 182

chosen and the covariance vector values are estimated using the semivariance 183

parameters and the model. A covariance matrix N x N is also estimated based on the 184

distance between each u and k position. The weight for each N position is estimated by 185

the product of the inverse of covariance matrix and the covariance vector. The residual 186

for the k position is estimated by the product of weight vector and the difference of z() 187

value for each N position and the general mean value. The estimated value for the k 188

position z(k) is the sum of residual value and the general mean value. Performing this 189

analysis varying the k value, the apparent density will be interpolated for the whole 190

stem and a consistent representation and its variation will be generated. 191

192

System of Equations 193

Model Development 194

We present a system of equations which integrate stem volume, wood density 195

and stem mass estimation. Assuming that the wood specific gravity is the ratio between 196

Page 8 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

9

the wood density and water density, and the water density is equal to 1 (g.cm-3 or Mg.m-197

3), we will use the expression (1) to estimate wood mass, in Mg. In this specific case, 198

following the x-ray methodology, the mass will be estimated with 12% of moisture 199

content, the volume will be green and the density will be treated as Apparent Wood 200

Density (AWD). 201

��� = � = ����(�)� ����(��) (1) 202

Individual stem volume can be estimated by using the solid of revolution 203

technique, namely, 204

�(������) = � ��[�(ℎ, � !,!")#]%ℎ&'()&'* (2) 205

Where E is the expected value and r is the radius (half diameter), in meters, 206

estimated for each h (height), in meters, for a stem with known diameter at breast height 207

(dbh) and total height (ht). The differential distance (%ℎ) is the value of the variation of 208

h (∆h) when ∆h goes to zero. The product of ��[�(ℎ, � !,!)#] is the area under the 209

stem profile curve and, multiplied by%ℎ, generates the volume. The relationship 210

between r and h has been well studies in forest research and has been modeled using 211

various taper functions. We used the following expression, derived from Kozak et al. 212

(1969), to estimate the expected value of squared radius: 213

�[�(ℎ, � !,!)#] = � !# ,-.* + -.0 &( + -.# 1&(2# + -.3 1&(234 (3) 214

The expression (3) could be any other taper function, either linear or nonlinear. 215

We have been using this one due to its simplicity and precision in estimating tree taper 216

(Table 2). 217

Substituting (3) into (2), we have: 218

Page 9 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

10

�(������) = � � 5� !# ,-.* + -.0 &( + -.# 1&(2# + -.3 1&(2346 %ℎ&'()&'* (4) 219

When the stem wood density value is known and there is no variation from base 220

to top and from the pith to the bark of the tree, the following relation would be 221

appropriate in order to estimate stem mass. 222

�(7899) = � ��[�(ℎ, � !,!")#]%ℎ&'&)&'* ∗ � (5) 223

If the density varies from base to top of the tree, we could use a function 224

E[�(ℎ)] instead of�, where the expression �[�(ℎ)] explains the expected density as a 225

function of height. 226

�(7899) = � ��[�(ℎ, � !,!")#]%ℎ&'&)&'* ∗ �[�(ℎ)] (6) 227

If the density varies from both the base to the top of the tree and pith to bark, the 228

variation would be explained by height and radius �[�(ℎ, �)] and the mass estimation 229

expression would be: 230

E(Mass)=∑ > � πE[�(ℎ, � !,!")#]δhh=ht@

h=0*E[ρ(h,r)]

-� πE A(�(ℎ, � !,!")-∆C)2D δh

h=ht@h=0

* EEρFh,r-∆CGHI0 by ∆Jr=rmax (7) 231

The expression (7) is a combination of continuous and discrete sum. The first 232

integral of the difference inside the braces estimates the stem mass with height varying 233

from base to the top of the tree and in radial position r. The second integral estimate the 234

stem mass, but with radius equal r-∆r, where ∆r is the incremental shell thickness. The 235

difference between the two integrals generates the shell mass, with expected density 236

E[ρ(h,r)]. The sum of the differences results in stem total mass. If we use the lim ∆r=0, 237

then the expression (7) would be: 238

Page 10 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

11

E(Mass)=� > � πE[�(ℎ, � !,!")2]δhh=ht@

h=0*E[ρ(h,r)]δr

-� πE[(�(ℎ, � !,!")-δr)2]δh

h=ht@h=0

*EEρFh,r-δrGHδrI0

r=rmax (8) 239

The differential distance (%�) is the value of the variation of r (∆r) when ∆r goes 240

to zero. The expression (8) will generate a model-unbiased estimate of the wood stem 241

mass. Accordingly, the unbiased estimation for the apparent wood density mean would 242

be: 243

E[ρ(h, �)] = N(����)N(� ����) (9) 244

The challenge here is to model both radial and longitudinal density variation. So, 245

with our densitometric data we chose to model (10) based on graphical relationship for 246

our database (Fig. 2). The nonlinear chosen function was the following concave upward 247

paraboloid relationship: 248

�[�(ℎ, �)] = 0OPQROPS(JT)ROPU1JT2U (10) 249

The expression (10) estimates the expected wood density from pith to bark (r) 250

and from base to top (h), having VW*, VW0, 8XYVW#the parameters to be estimated. 251

The functions (3) and (10) were fitted by R software (R Core Team, 2016), using 252

generalized least square (gls), generalized nonlinear least square (gnls) and nonlinear 253

mixed-efects model (nlme) libraries, respectively. 254

Model Performance and Evaluation 255

We adopted a generalized mixed-model approach which allowed us also to 256

address heteroscedasticity and autocorrelation. More details about this approach can be 257

found in Pinheiro and Bates (2000) and McCulloch and Searle (2001). 258

Page 11 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

12

The evaluation criteria used were Root Mean Square Error (RMSE) and 259

Coefficient of Determination (R2), presented in (11) and (12). After improving the 260

models by generalized mixed approach the Akaike Information Criterion (AIC) 261

(Sakamoto et al. 1986) and Likelihood Ratio Test (LRT) were used to evaluate the 262

model improvements. The expressions for these are shown in (13) and (14). 263

264

Z7[� = \∑ (]^_]W^)U`̂aS(b_c) (11) 265

Z# = ∑ (]dP _]e)U`̂aS∑ (]^_]e)U`̂aS ∗ 100 (12) 266

�hi = −2�(lm/o) + 2Xc�C (13) 267

pZq = 2ln(ptu#ptu0) In these formulas yi is the observed value for the ith observation, oWv is the 268

estimated value, n is the number of total observation, p is the number of parameters in 269

the model, oe is the mean of the observed values, �(lm/o) is the natural logarithm of the 270

profiled likelihood for the vector parameter lm as a function of the observed values for y, 271

Lik1 is the likelihood value for the restricted model and Lik2 is the likelihood value for 272

the general model. Under the null hypothesis that the restricted model is more adequate, 273

the distribution for LRT is χ2 with k2-k1 degrees of freedom. 274

275

Results 276

Fitting taper and apparent density functions 277

Page 12 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

13

Taper (3) and density (10) functions were fitted in order to model volume and 278

mass. Table 1 summarizes the statistics for the trees selected for taper and apparent 279

density measurements. The source of variation among plants for both taper and apparent 280

density databases are from genetic and environmental factors. Environmental factors 281

such as competition index, fertilization, pest and disease and seedling quality are 282

determinant source that explain the variation among the sampled trees. The upper stem 283

diameter measurements for the 72 sample trees, within three blocks, are displayed in 284

Fig. 1. As is apparent, the diameter at the base of trees varies from 15 to about 30 cm 285

and stem height varies from 6 to 15 meters. 286

Fig. 1. 287

The apparent density relationships with the ratio between radius, from pith to 288

bark, and height, from base to top, also including a loess smooth pattern, are displayed 289

in Fig. 2. The first remark is an observed decrease trend follows by an increase of the 290

apparent density when the ratio between radius and height increases. 291

Fig.2. 292

A better representation, including both radial and longitudinal density variation 293

for each tree, can be observed in Fig. 3, in which kriging geostatistical techniques were 294

applied to model the bidirectional variation. Also called Gaussian process regression, 295

with this technique it is possible to interpolate, or estimate, the density values in the 296

stem positions where the wood samples were not collected. As shown, the density, in 297

general, increases from pith to bark and from base to top.. 298

Parameter estimates and model statistics for taper function (3) and apparent 299

density function (10) are shown in Table 2. In both functions, the p-values for all 300

parameters were significant, considering α=0.05 (or 5%), ratifying the capacity of these 301

Page 13 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

14

models to explain the observed variation in stem volume and density within the stem. 302

When modeling stem taper, heteroscedasticity and autocorrelation was also modeled, 303

using the gls function from nlme R package (Pinheiro and Bates 2000). The Akaike 304

Information Criterion (AIC) decreases from -2188 to -3030, having a likelihood ratio 305

test value of 846 and p-value <0.0001, meaning that the heteroscedastic and 306

autocorrelated model had better performance than a reduced model assuming 307

homoscedastic and independent observations. 308

Fig. 3 309

Fig. 4 depicts, for taper equation, the improvement in the residual distribution 310

after modeling heteroscedasticity and autocorrelation. For apparent density function, 311

due to the height variation between trees and within each tree, heteroscedasticity and 312

autocorrelation were also modeled by considering the model as a two-level mixed-313

effect: tree level and height position level within tree level (Pinheiro and Bates 2000). 314

The AIC values before and after modeling were, respectively, -1701 and -2513, 315

generating a likelihood ratio test value of 828 and p-value <0.0001. Fig. 5 shows a 316

centered in zero and constant residual distribution within and among trees after 317

heteroscedasticity and autocorrelation modeling. A better resolution of the curvilinear 318

U-shape observed and estimated apparent density for tree 5 in the base position is 319

shown in Fig. 6. 320

321

Fig. 4. 322

323

Fig. 5. 324

325

Page 14 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

15

Fig. 6. 326

Estimating mass 327

Density varying from base to top 328

In order to exemplify the stem biomass assessment, the functions (3) and (10), 329

after fitting, were used to estimate the partial volume and mass of a sample tree, picked 330

among 72 harvested trees, with dbh inside bark of 25 cm and a total height of 11.65 m 331

(Table 3). The radius values, for each height position, were estimated by the taper 332

function. The density estimates were called marginal because its estimate used as 333

independent variables the height position and the external values of inside bark radius 334

for each height position. The apparent wood density, Mg.m-3, decreases from 0.4735, at 335

height of 0.15 m, to 0.3949, at height of 0.65 m. After that, the density increases until 336

the top of the tree, reaching 0.4238. The average density, stem volume and stem mass 337

were, respectively, 0.4190 Mg.m-3, 0.2734 m3 and 0.1147 Mg. A common practice is to 338

multiply the dbh density value (0.4016 Mg.m-3) with stem volume (0.2737 m3) in order 339

to estimate the stem mass. If we do so, the estimated mass is 0.1099 Mg and, compared 340

to 0.1147 Mg, which demonstrates a bias of 4.18%. 341

342

Density varying from base to top and from pith to bark. 343

Also for a 25 cm dbh and 11.65 m height tree, the fitted functions (3) and (10) 344

were substituted into the function (7), with ∆=2.5 cm, in order to generate a more 345

precise mass estimation. This procedure subdivided the tree into 4 shells and a core 346

stem. The shell thickness and the core stem radius at breast height were 2.5 cm, 347

increasing to the tree base and decreasing to the tree top, following the sigmoidal-shape 348

taper function estimates. The results are displayed in Table 4. The outer shell, from 20 349

Page 15 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

16

to 25 cm of dbh, represents roughly 40% of the total volume and mass. This percentage 350

decreases about 10% for each inner shell and for core stem. The estimated stem volume 351

was 0.2737 m3, which is the same value in Table 3, showing consistent estimates for 352

each shell and for stem core. The estimated stem mass was 0.1131 Mg. This value was 353

estimated by the sum of the mass for each shell and core stem. That is, base to top and 354

pith to bark density values were multiplied by each volume value to generate the stem 355

mass. Compared to the mass value estimated by taking the outer density from base to 356

top, 0.1147 Mg (Table 3), the bias is 1.4%. This value increases with the density 357

variation in both from base to top and from pith to bark. The mean density, estimated 358

by expression (9), was 0.4132 Mg.m-3. In the sample tree, this value is located at heights 359

around 2.5m, 2.0m, 1.5m, 1.0m, and 0.5m for the first, second, third, fourth shell and 360

core stem, respectively. The stem mass also was estimated using the double integral, 361

with limit of ∆r goes to 0, following expression (8). The estimated value was 0.1128 362

Mg, which is very close to the value estimated by summing, with ∆r=2.5 cm (0.1131 363

Mg), and the bias was 0.26%. Taking in account the stem volume, 0.2737 cubic meters, 364

the mean density was 0.4121 Mg.m-3, which is a slightly smaller than the estimated 365

value in expression (7). 366

367

Discussion 368

Taper and density functions 369

The relatively small variation in dbh and height in our sample trees is due to the 370

young plantation age, about 4.7 years old. In apparent density database, the minimum 371

threshold dbh was fixed at 15 cm, since logs will be our final gross product. The greater 372

vertical and horizontal variation of the observed apparent density values are due to 373

Page 16 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

17

sensitivity and precision of the x-ray method, which captured every irregularity inside 374

of stem wood, like knots and cracks, increasing the variability of the density values. 375

Also, the method recorded some intermittent peaks in the observed apparent density that 376

could be related to the wood rings: the lower peaks represent early wood and the higher 377

peaks late wood. Studying a 50-year-old black spruce plantation, Koubaa et al. (2002) 378

found a variation in density of 0.3 (early wood) to 0.66 (late wood) in the twentieth ring. 379

The apparent density values increased from pit to bark and from base to top and 380

approximately the same behavior was found by Jyske et al., (2008) studying Norway 381

Spruce stems. Zobel and van Buijtenen (1989) state that the wood density variation 382

within the tree stem is different among species. Nock, et al. (2009), also using X-Ray 383

technology, found that wood density for Toona ciliata, in Thailand, increased 27% from 384

pith to bark 385

The polynomial taper function (3) used in this study, after Kozak (1969), has 386

been also fitted by other researchers and it almost always has been found to be very 387

precise. This function is very flexible in modeling the sigmoidal stem shape, in both 388

plantation and natural situation. Although the nonlinear apparent density function (10) 389

has not been applied in other previous studies, it modeled very well the U-shape density 390

variation as a function of the ratio between stem radius and height position. The 391

performance for both functions was improved by modeling heteroscedasticity and 392

autocorrelation. The apparent density function also was modeled by multilevel mixed-393

effect model approach, accessing the variations between and within tree, which 394

improved the estimating process. This will influence the mass estimation performance. 395

The apparent density estimated value of 0,419 Mg.m-3 is consistent with 0,37 Mg.m-3 396

found by Trianoski et al. (2011), using red cedar trees, from plantation and with 18 397

Page 17 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

18

years old. This value would be close to 0.4 Mg.m-3 in apparent density, instead specific 398

gravity, due to 12% of humidity. 399

400

Mass estimation 401

When the stem mass was estimated taking in account the density variation from 402

base to the top of the tree, the average estimated density was found at approximately 5.0 403

meters of height, about 43% of the total height. Pérez-Cruzado and Rodríguez-Soalleiro 404

(2009), studying Eucalyptus nittens, found that the average basic density occurs at a 405

relative height of 30–35% along the stem. Compared to the mass estimated by using the 406

apparent density value at dbh, the bias generated was 4,2%. This is a significant value 407

compared to, for example, forest sample error, which could be, in some situation, 408

smaller than 5%. Indeed, the bias will be directly related to the variation of the base-to-409

top apparent density and when the apparent density mean value is located far from dbh 410

position. Greater variation, in this case, implies greater bias. 411

When stem mass was estimated considering the density variations from base 412

to top and form pith to bark, considering the radius variation as a discrete approach 413

(∆=2.5 cm), the bias was 1,4%, compared to the mass estimated accounted just with 414

base to top density variation. Again, the bias would be correlated to the stem density 415

variation, as opposed to assuming constant density presumably from its value at breast 416

height. 417

418

Conclusions 419

In general, the apparent density values increased from base to top and from the 420

pith to the bark for each tree, in a nonlinear U-shape trend. The proposed system of 421

Page 18 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

19

equations fitted very well the tree taper and the tree density variation. Compared to the 422

traditional method, which the density values at dbh are used to estimate the stem mass, 423

the bias estimation was reduced by accessing the density variation from the base to the 424

top of the tree. Also, the estimation of mass was improved by modeling the density 425

variation from both base to top and pith to bark. The model performance was improved 426

by using the generalized linear and nonlinear mixed-effects approach. The system can 427

be used to estimate stem volume, density and mass for every red cedar tree a least 5 428

years old, at the study location, having dbh and total height measurements. As a future 429

study suggestion, using older tree and different sites database would generate a more 430

practical application system for this and other plant species. 431

432

Acknowledgements 433

The authors thank to CNPq and Fapemig, the Brazilian and Minas Gerais State research 434

sponsors, respectively. 435

436

References 437

Behre, C.E. 1927. Form-class taper curves and volume tables and their application. 438

Journal of Agricultural Research 35(8): 673-744. 439

Chidester G.H., McGovern I.N., McNaughton G.C. 1938. Comparison of sulfite pulps 440

from fast grown loblolly, shortleaf, longleaf and slash pines. Paper Tr J 107, 441

Tappi Sec: 32-35 442

Clutter, J.L. 1980. Development of taper functions from variable-top merchantable 443

volume equations. Forest Science 26(1):117-120. 444

Page 19 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

20

Demaerschalk, J.P. 1973. Integrated systems for the estimation of tree taper and 445

volume. Canadian Journal of Forest Research. 3(90):90-94. 446

Downes G.M., Hudson I.L., Raymond C.A., Dean G.H., Michell A.J., Schimleck L.R., 447

Evans R., Muneri A. 1997. Sampling plantation eucalypts for wood and fibre 448

properties, CSIRO Publishing, Melbourne, 1997. 449

Edmonds, J. 1993. The potential value of Toona species (Meliaceae) as multipurpose 450

and plantation trees in Southeast Asia. The Commonwealth 451

ForestryReview, 72(3), 181-186. Retrieved from 452

http://www.jstor.org/stable/42616713 453

Frederick, D. J., Madgwick H. A. I., Oliver, G. R. 1982. Wood basic density and 454

moisture content of young Eucalyptus regnans grown in New Zealand. New 455

Zealand Journal of Forestry Science 12(3): 49. 456

Furnival, George M. 1961. An index for comparing equations used in constructing 457

volume tables. Forest Science. 7: 337-341. 458

Gregoire, T.G. and O. Schabenberger. 1996. A Non-linear mixed-effects model to 459

predict cumulative bole volume of standing trees. Journal of Applied Statistics 460

23(2&3):257-271. 461

Gregoire, T.G. ,Schabenberger, O., Kong F. 2000. Prediction from an Integrated 462

Regression Equation: A Forestry Application. Biometrics 56. 414-419. 463

Heitz, P., Valencia, R., Wright, S. J. 2013. Strong radial variation in wood density 464

follows a uniform pattern in two neotropical rain forests. Functional Ecology. 465

Höjer, A. G. 1903. Bihang till fr. loven: Om vӓra barrskogar. Stockholm. 466

Hua P., Edmonds J.M. 2008. Toona (Endlicher) M. Roemer, Fam. Nat. Syn. Monogr. 1: 467

131. 1846. In: Flora of China, vol. 11: Oxalidaceae through Aceraceae. St Louis, 468

MO, USA: Missouri Botanical Garden, 112-115. 469

Page 20 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

21

INMET. 2016. Climatological station net. Brazilian Meteorological Institute. (accessed 470

in 07/05/2016 - http://www.inmet.gov.br). 471

Jonson, T.1910. Taxatoriska undersökningar om skogsträdens form. I. Granens 472

Stamform. Skogsvardsför. Tidskr. 8 (11): 285-328, illus. 473

Jyske, T., Mäkinen, H., Saranpää, P. 2008. Wood density within norway spruce stems. 474

Silva Fennica 42(3) 475

Köppen, W. 1900. Versuch einer klassifikation der klimate, vorzugsweise nach ihren 476

Beziehungen zur Pflanzenwelt. Geographische Zeitschrift 6, 657–679. 477

Koubaa A., Zhangb, S.Y.T. and Maknic S. 2002. Defining the transition from 478

earlywood to latewood in black spruce based on intra-ring wood density profiles 479

from X-ray densitometry. Ann. For. Sci. 59 511–518. 480

Kozak, A., D.D. Munro, and J.H.G. Smith. 1969. Taper functions and their application 481

in forest inventory. The Forestry Chronicle 45:278-283. 482

Krige, Danie G. 1951. A statistical approach to some basic mine valuation problems on 483

the Witwatersrand. J. of the Chem., Metal. and Mining Soc. of South 484

Africa. 52 (6): 119–139. 485

Luxford, R. F., Markwardt, L.J. 1932. The strength and related properties of redwood. 486

USDA Technical Bulletin No. 305. Washington, D.C. 487

Matheron, G. 1962. Traité de géostatistique appliquée. Editions Technip. 488

489

McCulloch, C.E. and Searle, S.R. 2001. Generalized, Linear, and Mixed Models, 490

Wiley, New York. 491

Nock, C.A., Geihofer, D., Grabner M., Baker,P.J., Bunyavejchewin S. and Hietz, P. 492

2009. Wood density and its radial variation in six canopy tree species differing in 493

shade-tolerance in western Thailand. Annals of Botany 118:1-10. 494

Page 21 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

22

Parresol, B.R., and Thomas, C.E. 1989. A density-integral approach to estimating stem 495

biomass. For. Ecol. Manage. 26: 285-297. 496

Pérez-Cruzado C. and Rodríguez-Soalleiro R. 2009. Improvement in accuracy of 497

aboveground biomass estimation in Eucalyptus nitens plantations: Effect of bole 498

sampling intensity and explanatory variables. Forest Ecology and Management. 499

261. 2016-2028. 500

QMS (QUINTEK MEASUREMENT SYSTEMS). 1999. Tree ring analyzer users guide 501

- Model QTRS-01X. Knoxville.72 p. 502

Quilhó, T., Pereira, H. 2001. Within and between tree variation of bark content and 503

wood density of Eucalyptus globulus in commercial plantations. Iawa Journal, 504

Vol. 22 (3), 2001: 255–265. 505

Raymond, C.A. and Joe, B. 2007. Patterns of basic density variation for Pinus 506

radiata grown in south-west slopes region of New South Wales, Australia. New 507

Zealand Journal of Forestry Science 37(1): 81–95. 508

Ribeiro Jr., P.J. & Diggle, P.J. 2001. geoR: A package for geostatistical analysis. R-509

NEWS, Vol 1, No 2, 15-18. ISSN 1609-3631. 510

Romagnoli, M., Cavalli, D., Spina, S. 2014. Chestnut wood quality. BioResources 9(1). 511

R Core Team (2016). R: A language and environment for statistical computing. R 512

Foundation for Statistical Computing, Vienna, Austria. URL https://www.R-513

project.org/. 514

Rueda R., Williamson G. B. 1992. Radial and vertical wood specific gravity 515

in Ochroma pyramidale (Cav. ex Lam.) Urb. (Bombacaceae). Biotropica 24:512–516

518. 517

Sakamoto, Y., Ishiguro, M. and Kitagawa, G. 1986. Akaike Information Criterion 518

Statistics, Reidel, Dordrecht, Holland. 519

Page 22 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

23

Schiffel, A. 1905. Form und inhalt der lärche. Mitt. Forstl. Versuchsw. Österr., Heft 31, 520

122 p., illus. 521

Schumacher, F.X. 1926. A Method of measuring form quotient of standing trees. 522

Journal of Forestry 24:552-554. 523

Trianoski, R., Iwakiri, S., De Matos, J.L.M. 2011. Potential use of planted fast-growing 524

species for production of particleboard. Journal of Tropical Forest Science, 311-525

317. 526

Vader, J. 1987. Red cedar: the tree of Australia’s history. Reed. NSW. pp. 200. 527

Wiemann, M. C., and Williamson,G.B. 1989a. Radial gradients in the specific gravity 528

of wood in some tropical and temperate trees. Forest Science 35 : 197 – 210 . 529

Zobel, B.J., van Buijtenen, J. P.1989 . Wood variation: Its causes and control. Spring 530

Verlag, New York. 531

Page 23 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

Fig. 1. Upper stem diameter measurements for 72 trees in 3 experimental blocks.

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

oo

o

o

o

o

o

o

o

o

o

o

o

o

oo

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

0 5 10 15 20 25 30

05

10

15

diameter inner bark(cm)

Tre

e h

eig

ht (b

ase to top)

(m

)

Block 1

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

0 5 10 15 20 25 30

05

10

15

diameter inner bark(cm)

Block 2

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

0 5 10 15 20 25 30

05

10

15

diameter inner bark(cm)

Block 3

Page 24 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

Fig.2. Variation of the apparent density as a function of the ratio between radius and

height for 6 sample trees with a loess smoother line to portray the pattern of the

relationship.

0.3

0.4

0.5

0.6

0.7

0.8

0 20 40 60

Tree 1 Tree 2

0 20 40 60

Tree 3

Tree 4

0 20 40 60

Tree 5

0.3

0.4

0.5

0.6

0.7

0.8Tree 6

Radius(cm)/Height(m)

Appare

nt

Density(M

g/m

3)

Page 25 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

Figure 3. Kriging of observed apparent density for sample trees

Page 26 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

Fig. 4. Residual distribution before (A) and after (B) heteroscedasticity and

autocorrelation modeling for taper model.

(A)

Fitted Taper Values (r/dbh)²)

Sta

ndard

ized r

esid

uals

-4

-2

0

2

4

0.0 0.1 0.2 0.3 0.4

Ѫ

ѪѪ

ѪѪ

ѪѪѪ

ѪѪ

Ѫ

Ѫ

Ѫ

ѪѪ

Ѫ

Ѫ

Ѫ

Ѫ

ѪѪ

ѪѪѪ

Ѫ

ѪѪ

ѪѪ

ѪѪ

Ѫ

Ѫ

Ѫ

Ѫ

ѪѪ

ѪѪѪ

ѪѪ

ѪѪ

ѪѪѪѪ

Ѫ

Ѫ

Ѫ

Ѫ

ѪѪѪ

ѪѪѪ

ѪѪ

ѪѪ

Ѫ

ѪѪ

ѪѪѪ

ѪѪѪ

Ѫ

ѪѪѪ

ѪѪ

ѪѪ

ѪѪ

Ѫ

Ѫ

ѪѪѪ

Ѫ

ѪѪ

ѪѪѪѪ

ѪѪ

ѪѪ

ѪѪѪѪ

Ѫ

Ѫ

ѪѪ

Ѫ

Ѫ

ѪѪѪ

Ѫ

ѪѪ

ѪѪѪѪ

Ѫ Ѫ

ѪѪ

Ѫ

ѪѪѪѪ

Ѫ

ѪѪѪ

Ѫ

ѪѪѪ

Ѫ

Ѫ

Ѫ

ѪѪѪѪ

Ѫ Ѫ

Ѫ

Ѫ

Ѫ

ѪѪ

ѪѪ

Ѫ

ѪѪ

ѪѪ

ѪѪ

Ѫ

Ѫ

ѪѪ

ѪѪѪѪ

Ѫ

Ѫ

ѪѪ

ѪѪ

ѪѪѪ

ѪѪ

Ѫ

ѪѪ

ѪѪѪ

Ѫ

ѪѪ

ѪѪ

ѪѪ

Ѫ

Ѫ

ѪѪѪѪѪ

Ѫ

Ѫ

ѪѪ

ѪѪ

ѪѪ

ѪѪ

Ѫ

Ѫ

ѪѪ

ѪѪѪ

Ѫ

Ѫ

ѪѪ

Ѫ

ѪѪ

ѪѪ

ѪѪ

ѪѪ

ѪѪ

ѪѪ

Ѫ

ѪѪ

ѪѪ

Ѫ ѪѪѪ

ѪѪ

Ѫ

ѪѪ

Ѫ

Ѫ

ѪѪ

ѪѪѪѪ

Ѫ

ѪѪ

ѪѪ

ѪѪ

Ѫ

Ѫ

Ѫ

Ѫ

ѪѪ

ѪѪ

Ѫ

Ѫ

ѪѪ

Ѫ

ѪѪѪ

Ѫ

Ѫ

ѪѪ

ѪѪ

ѪѪѪ

Ѫ

ѪѪ

ѪѪ

ѪѪ

Ѫ

Ѫ

ѪѪ

ѪѪ

ѪѪ

Ѫ

ѪѪѪ

ѪѪѪѪ

Ѫ

ѪѪѪ

ѪѪ

ѪѪѪ

Ѫ

ѪѪ

ѪѪѪ

ѪѪ

Ѫ

ѪѪ

ѪѪ

ѪѪ

Ѫ

Ѫ

Ѫ

ѪѪ

Ѫ

ѪѪ

Ѫ

ѪѪ

ѪѪѪ

ѪѪ Ѫ

ѪѪ

ѪѪ

ѪѪѪ

Ѫ

ѪѪѪ

ѪѪѪ

Ѫ

Ѫ

ѪѪ

ѪѪѪ

ѪѪ

ѪѪ

Ѫ

ѪѪ

ѪѪѪ

Ѫ

ѪѪ

ѪѪѪѪѪ

Ѫ

Ѫ

ѪѪѪѪ

Ѫ

ѪѪ

ѪѪ

ѪѪ

ѪѪ

Ѫ ѪѪ

Ѫ

ѪѪ

ѪѪѪ

ѪѪ

Ѫ

ѪѪѪѪ

Ѫ

Ѫ

ѪѪѪѪ

ѪѪ

Ѫ

Ѫ

Ѫ

ѪѪ

Ѫ

ѪѪ

Ѫ

Ѫ

ѪѪ

ѪѪ

Ѫ

ѪѪ

Ѫ

ѪѪ

Ѫ

ѪѪѪѪ

Ѫ

ѪѪ

ѪѪ

ѪѪѪ

Ѫ

Ѫ

Ѫ

ѪѪ

ѪѪѪ

Ѫ

ѪѪѪ

ѪѪ

ѪѪ

ѪѪ

ѪѪ

ѪѪѪ

ѪѪѪ

ѪѪѪѪѪ

Ѫ

Ѫ

Ѫ

Ѫ

ѪѪ

ѪѪ

ѪѪѪ

Ѫ

Ѫ

ѪѪѪ

Ѫ

ѪѪѪѪ

ѪѪѪ

ѪѪѪ

Ѫ

Ѫ

Ѫ

Ѫ

ѪѪѪѪ

ѪѪ

ѪѪ

Ѫ

Ѫ

ѪѪ

ѪѪ

ѪѪѪ

ѪѪѪ

ѪѪѪѪ

Ѫ ѪѪѪ

ѪѪ

ѪѪѪ

ѪѪ

Ѫ

ѪѪ

ѪѪ

Ѫ

(B)

Fitted Taper Values (r/dbh)²

Sta

ndard

ized r

esid

uals

-4

-2

0

2

4

0.0 0.1 0.2 0.3 0.4

Ѫ

ѪѪѪ

Ѫ

ѪѪ

ѪѪ

ѪѪѪ

ѪѪ

ѪѪ

Ѫ

ѪѪѪ

ѪѪѪ

ѪѪ

ѪѪ

ѪѪ

ѪѪ

Ѫ

Ѫ

Ѫ

ѪѪѪ

ѪѪѪ

ѪѪѪѪ

Ѫ

Ѫ

Ѫ Ѫ

ѪѪѪ

Ѫ

ѪѪ

Ѫ

ѪѪѪѪ

Ѫ

Ѫ

Ѫ

Ѫ

ѪѪѪѪѪ

Ѫ

ѪѪ

Ѫ

ѪѪѪ

ѪѪ

ѪѪ

ѪѪ

Ѫ

Ѫ

Ѫ

Ѫ

ѪѪ

ѪѪѪ

Ѫ

Ѫ

Ѫ

Ѫ Ѫ

ѪѪѪѪѪ

ѪѪ

Ѫ

ѪѪѪ

ѪѪѪ

Ѫ

Ѫ

ѪѪѪ

ѪѪѪ

Ѫ Ѫ

ѪѪѪ

Ѫ

ѪѪѪ

Ѫ

ѪѪѪ

Ѫ

ѪѪ

Ѫ

Ѫ

Ѫ

ѪѪ

Ѫ

ѪѪ

Ѫ ѪѪ

ѪѪ

Ѫ

Ѫ

Ѫ

Ѫ

ѪѪѪ

ѪѪ

Ѫ

Ѫ

Ѫ

Ѫ

ѪѪ

ѪѪ

Ѫ

ѪѪ

Ѫ

ѪѪѪ

ѪѪѪ

Ѫ

ѪѪ

ѪѪѪ

ѪѪ

Ѫ Ѫ

ѪѪ

ѪѪ

Ѫ

ѪѪ

Ѫ

ѪѪѪ

Ѫ

ѪѪ

Ѫ

ѪѪ

ѪѪ

Ѫ

ѪѪ Ѫ

ѪѪ

Ѫ

Ѫ

Ѫ

ѪѪ Ѫ

ѪѪ

Ѫ

Ѫ

Ѫ

Ѫ

Ѫ Ѫ

ѪѪ

Ѫ

ѪѪ

Ѫ

ѪѪѪ

ѪѪѪ

Ѫ

Ѫ ѪѪѪ

Ѫ

ѪѪ

Ѫ

Ѫ

Ѫ

Ѫ

Ѫ

ѪѪ

Ѫ

ѪѪ

Ѫ

ѪѪ

ѪѪ

Ѫ

Ѫ

Ѫ

Ѫ

Ѫ

ѪѪ

ѪѪ

Ѫ

Ѫ

ѪѪѪ

Ѫ

ѪѪѪ

ѪѪ

ѪѪѪ

Ѫ

ѪѪ

Ѫ

Ѫ

ѪѪѪ

Ѫ

Ѫ

Ѫ

Ѫ

Ѫ

ѪѪ

ѪѪ

Ѫ

Ѫ

ѪѪѪѪ

ѪѪѪ

Ѫ

ѪѪѪ

ѪѪѪ

ѪѪ

Ѫ

Ѫ

ѪѪ

ѪѪ

Ѫ

Ѫ

Ѫ

Ѫ

ѪѪ

ѪѪ

Ѫ

Ѫ

Ѫ

Ѫ

ѪѪ

Ѫ

Ѫ

Ѫ

Ѫ

Ѫ

ѪѪ

ѪѪ

Ѫ

ѪѪ Ѫ

ѪѪ

ѪѪѪ

Ѫ

ѪѪ

ѪѪѪ

ѪѪѪ

Ѫ

Ѫ

ѪѪѪ

Ѫ

Ѫ

Ѫ

Ѫ

ѪѪѪѪ

ѪѪѪ

Ѫ

Ѫ

ѪѪ

ѪѪ

Ѫ

Ѫ

Ѫ

Ѫ

ѪѪѪ

Ѫ

Ѫ

Ѫ

ѪѪ

ѪѪ

Ѫ

Ѫ

ѪѪ

Ѫ ѪѪ

ѪѪѪ

ѪѪ

Ѫ ѪѪѪѪѪ

Ѫ

Ѫ

ѪѪ

ѪѪѪѪ

ѪѪ

Ѫ

Ѫ

Ѫ

ѪѪ

Ѫ

Ѫ

Ѫ

ѪѪ

ѪѪ

ѪѪ

Ѫ

Ѫ

ѪѪ

ѪѪ

ѪѪ

ѪѪ

ѪѪ

ѪѪѪ

Ѫ

ѪѪ

Ѫ

Ѫ

Ѫ

Ѫ

Ѫ

ѪѪ

ѪѪ

Ѫ

ѪѪѪ

Ѫ

Ѫ

ѪѪѪ

ѪѪѪ

ѪѪ

ѪѪ

ѪѪѪѪѪ

Ѫ

Ѫ

Ѫ

Ѫ

Ѫ

ѪѪ

Ѫ

ѪѪ Ѫ

ѪѪ

Ѫ

Ѫ

Ѫ

ѪѪ

Ѫ

ѪѪѪ

ѪѪ

ѪѪ

ѪѪѪ

ѪѪ

Ѫ

Ѫ

ѪѪѪ

Ѫ

ѪѪ

ѪѪѪ

Ѫ

ѪѪ

Ѫ

ѪѪ

ѪѪѪ

ѪѪѪѪ

Ѫ

ѪѪ Ѫ

ѪѪѪ

ѪѪ

ѪѪ

ѪѪ

Ѫ

ѪѪ

ѪѪ

Ѫ

Page 27 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

-3

-2

-1

0

1

2

3

0.40 0.45 0.50 0.55 0.60

Tree 1 Tree 2

0.40 0.45 0.50 0.55 0.60

Tree 3

Tree 4

0.40 0.45 0.50 0.55 0.60

Tree 5

-3

-2

-1

0

1

2

3

Tree 6

Apparent Density Fitted Values (Mg/m3)

Sta

ndard

ized r

esid

uals

Fig. 5. Standardized residuals versus fitted apparent density by tree.

Page 28 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

Fig. 6. Observed and estimated values for tree 5 in the base position.

Radius(cm) - From Pith to Bark

Appare

nt D

ensity(M

g/m

³)

0.2

0.3

0.4

0.5

0.6

0.7

2 4 6 8 10

Tree5/Base

Page 29 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

Table 1 – Descriptive statistics for 72 trees used on fitting taper function and 6 trees for

measuring x-ray apparent wood density.

Statistics

Variables for Taper Variables for Apparent Density

DBH

(cm)

Height

(m)

Volume

(m3)

DBH

(cm)

Height

(m)

Volume

(m3)

Minimum 8.8 6.3 0.02 15.0 9.1 0.06

Average 18.3 10.7 0.09 17.7 10.9 0.09

Maximum 25.4 13.6 0.15 22.8 13.6 0.15

Standard

Deviation 3.0 1.6 0.03 2.8 1.7 0.03

Page 30 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

Table 2. Estimated parameters and its statistics for taper and density function.

Taper Function

Parameter Estimated

Value

Standard

Error t - value Pr(>|t|)

β0 0.38425 0.00627 61.3 <0.0001

β1 -0,82719 0.03448 -23.9 <0.0001

β2 0,546438 0.05882 9.3 <0.0001

β3 0,1036911 0.029617 -3.5 0.0005

RMSE

(cm.cm-2

) 0.0591 - - -

R2(%) 94.18 - - -

Apparent Density Function

α0 2.30498 0.099096 23.26 <0.0001

α1 0.01898 0.006011 3.15 0.0160

α2 -0.000443 0.000088 -5.04 <0.0001

RMSE

(Mg.m-3

) 0.0888 - - -

Page 31 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

Table 3. Estimates for radius, density, volume and mass for a 25 dbh inside bark tree.

Height

(m)

Estimated

Radius o.b.

(cm)

Estimated

Marginal

Density

(Mg.m-3

)

Partial

Volume

Estimated

(m3)

Estimated

Mass (Mg)

0,15 15.4 0.4735 0.0352 0.01665

0.65 14.5 0.3949 0.0309 0.01222

1.15 13.6 0.3998 0.0085 0.00341

1.30 13.3 0.4016 0.0186 0.00748

1.65 12.7 0.4058 0.0238 0.00965

2.15 11.9 0.4099 0.0208 0.00852

2.65 11.1 0.4128 0.0182 0.00750

3.15 10.4 0.4149 0.0159 0.00658

3.65 9.7 0.4165 0.0139 0.00577

4.15 9.1 0.4177 0.0121 0.00506

4.65 8.5 0.4186 0.0106 0.00445

5.15 8.0 0.4194 0.0093 0.00392

5.65 7.5 0.4200 0.0083 0.00347

6.15 7.0 0.4205 0.0073 0.00307

6.65 6.6 0.4209 0.0065 0.00273

7.15 6.2 0.4213 0.0058 0.00243

7.65 5.9 0.4216 0.0051 0.00216

8.15 5.5 0.4218 0.0045 0.00190

8.65 5.2 0.4221 0.0039 0.00164

9.15 4.8 0.4223 0.0033 0.00138

9.65 4.3 0.4225 0.0026 0.00109

10.15 3.8 0.4227 0.0018 0.00078

10.65 3.0 0.4230 0.0010 0.00042

11.15 1.8 0.4233 0.0005 0.00020

11.65 0.0 0.4238 0.0000 0.00000

Totals 0.2737 0.11470

Means 0.4190 - -

Page 32 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research

Draft

Table 4. Volume and mass estimated for each shell for a 25 dbh inside bark tree.

Volume

(m3)

Mass

(Mg)

Volume

(m3)

Mass

(Mg)

Volume

(m3)

Mass

(Mg)

Volume

(m3)

Mass

(Mg)

Volume

(m3)

Mass

(Mg)

Volume

(m3)

Mass

(Mg)

0,15 0,0128 0,0061 0,0099 0,0048 0,0071 0,0038 0,0042 0,0018 0,0011 0,0004 0,0352 0,0169

0,65 0,0115 0,0045 0,0089 0,0035 0,0062 0,0025 0,0036 0,0015 0,0007 0,0003 0,0309 0,0123

1,15 0,0102 0,0041 0,0079 0,0032 0,0055 0,0022 0,0031 0,0013 0,0005 0,0002 0,0272 0,0110

1,65 0,0091 0,0037 0,0070 0,0028 0,0048 0,0020 0,0026 0,0011 0,0003 0,0001 0,0238 0,0097

2,15 0,0081 0,0033 0,0062 0,0025 0,0042 0,0017 0,0021 0,0009 0,0002 0,0001 0,0208 0,0086

2,65 0,0072 0,0030 0,0054 0,0023 0,0036 0,0015 0,0018 0,0007 0,0001 0,0001 0,0182 0,0075

3,15 0,0064 0,0027 0,0048 0,0020 0,0031 0,0013 0,0015 0,0006 0,0000 0,0000 0,0159 0,0066

3,65 0,0057 0,0024 0,0042 0,0018 0,0027 0,0011 0,0013 0,0005 0,0139 0,0058

4,15 0,0050 0,0021 0,0037 0,0015 0,0024 0,0010 0,0011 0,0004 0,0121 0,0051

4,65 0,0044 0,0019 0,0033 0,0014 0,0021 0,0009 0,0009 0,0004 0,0106 0,0045

5,15 0,0039 0,0017 0,0029 0,0012 0,0018 0,0008 0,0007 0,0003 0,0093 0,0039

5,65 0,0035 0,0015 0,0025 0,0011 0,0016 0,0007 0,0006 0,0002 0,0083 0,0035

6,15 0,0031 0,0013 0,0023 0,0010 0,0015 0,0006 0,0004 0,0002 0,0073 0,0031

6,65 0,0028 0,0012 0,0020 0,0009 0,0014 0,0006 0,0002 0,0001 0,0065 0,0027

7,15 0,0025 0,0011 0,0019 0,0008 0,0013 0,0006 0,0001 0,0000 0,0058 0,0024

7,65 0,0023 0,0010 0,0017 0,0007 0,0011 0,0005 0,0051 0,0022

8,15 0,0021 0,0009 0,0016 0,0007 0,0008 0,0003 0,0045 0,0019

8,65 0,0019 0,0008 0,0015 0,0006 0,0004 0,0002 0,0039 0,0016

9,15 0,0018 0,0008 0,0014 0,0006 0,0001 0,0000 0,0033 0,0014

9,65 0,0017 0,0007 0,0009 0,0004 0,0026 0,0011

10,15 0,0015 0,0006 0,0003 0,0001 0,0018 0,0008

10,65 0,0010 0,0004 0,0010 0,0004

11,15 0,0003 0,0001 0,0003 0,0001

11,65 0,0000 0,0000 0,0000 0,0000

Totals 0,1091 0,0457 0,0802 0,0338 0,0517 0,0223 0,0240 0,0100 0,0031 0,0013 0,2737 0,1131

TotalsShell 1

(20-25 cm dbh)

Shell 2

(15-20 cm dbh)

Shell 3

(10-15 cm dbh)

Shell 4

(5-10 cm dbh)

Core Stem

(5 cm dbh)Height

(m)

Page 33 of 33

https://mc06.manuscriptcentral.com/cjfr-pubs

Canadian Journal of Forest Research