Forest Ecology and Management 479 (2021) 118624

Contents lists available at ScienceDirect

Forest Ecology and Management

journal homepage: www.elsevier.com/locate/foreco

Assessing the impact of fine-scale structure on predicting wood fibre T

attributes of boreal conifer trees and forest plots
Jean-François Côtéa, , Joan E. Lutherb, Patrick Lenzc, Richard A. Fournierd, Olivier R. van Liere
⁎

a
Natural Resources Canada, Canadian Forest Service – Canadian Wood Fibre Centre, Québec, QC G1V 4C7, Canada
b
Natural Resources Canada, Canadian Forest Service – Atlantic Forestry Centre, Corner Brook, NL A2H 6J3, Canada
c
Natural Resources Canada, Canadian Forest Service – Canadian Wood Fibre Centre, Québec, QC G1V 4C7, Canada
d
Department of Applied Geomatics, Centre d’Applications et de Recherche en TELédétection, Université de Sherbrooke, Sherbrooke, QC J1K 2R1, Canada
e
Natural Resources Canada, Canadian Forest Service – Canadian Wood Fibre Centre, Corner Brook, NL A2H 6J3, Canada

ARTICLE INFO ABSTRACT

Keywords: Information about wood fibre attributes (WFA) is important for optimizing forest resource management and
Wood fibre attribute increasing the competitiveness of the sector. Many factors influence WFA at both the plot (e.g., age, stand
3D architectural model density, climate, and disturbance) and tree (e.g., crown development, stem shape, branchiness) levels. Recently,
Terrestrial LiDAR the use of terrestrial lidar (t-lidar) systems in forest inventory has enabled the measurement of forest structural
Tree structure
attributes, which were almost impossible to acquire with traditional field measurements. Using t-lidar scans of
Plot structure
individual trees and the architectural model L-Architect, we reconstructed the structure of trees and plots
comprising balsam fir and black spruce in insular Newfoundland, Canada. Core samples extracted from con­
comitant trees were analyzed for a series of nine WFA. The impact of fine-scale structure on predictive models of
WFA was assessed with parametric and non-parametric approaches. A variable importance analysis demon­
strated that structural attributes derived from L-Architect describing the tree crown geometry, branching
structure, stem form, spatial competition and canopy material distribution were highly important in the resulting
models. The cross-validated percentage of variance explained for the WFA predictive models ranged from
12–56% and 5–80% at tree- and plot-levels respectively. The addition of fine-scale structure improved the
models by 10–31% and 0–53% when compared to models developed using only in situ measurements at tree-
and plot-levels respectively. Information on species (at tree level) and composition (at plot level) did not im­
prove the predictive capability of models developed with L-Architect fine-scale structure. The results indicate that
better characterisation of forest structure using t-lidar and an architectural model can lead to improved WFA
prediction and their combination opens opportunities to significantly enhance forest inventory.

1. Introduction
Forest end-users require information about stand, plot and in­
dividual tree characteristics, as well as intrinsic wood properties, to
support decisions along the forest value chain. Tree species composi­
tion, stand age, and site index are commonly used to predict the po­
tential quality and quantity of timber available to serve various market
demands. The economic value of saw logs is largely affected by the
quality of the wood fibre, log shape and curvature, and number and
diameter of branches; whereas the value of pulpwood is largely de­
termined by wood fibre length, strength, and dimensions (Downes
et al., 2002). At the regional level, climate, topography, and soils have a
direct impact on the physical and chemical attributes of timber and
wood properties. The geographic variation at the regional level can lead
to different growing conditions for forest stands with impacts on nu­
merous factors including species composition, fertility class, and silvi­
cultural treatment. These large-scale factors govern the establishment
of different plant species that favour specific site conditions, and reg­
ulate light regime and photosynthesis, which in turn affect growth
(Väisänen et al., 1989; Smith and Hinckley, 1995; Wilhelmsson et al.,
2002). These alterations in tree growth might result in observable
variations in tree stem and crown morphology.
At the landscape and plot levels, wood fibre properties vary ac­
cording to stand development. Factors such as local topography, wind

Corresponding author at: Canadian Wood Fibre Centre, Natural Resources Canada / Government of Canada, 1055 Du P.E.P.S. Street, P.O. Box 10380, Québec,
⁎

Québec, G1V 4C7, Canada.

E-mail addresses: jean-francois.cote@canada.ca (J.-F. Côté), joane.luther@canada.ca (J.E. Luther), patrick.lenz@canada.ca (P. Lenz),
richard.fournier@usherbrooke.ca (R.A. Fournier), olivier.vanlier@canada.ca (O.R. van Lier).

https://doi.org/10.1016/j.foreco.2020.118624
Received 15 May 2020; Received in revised form 14 September 2020; Accepted 15 September 2020
Available online 30 September 2020
0378-1127/ Crown Copyright © 2020 Published by Elsevier B.V. This is an open access article under the CC BY-NC-ND license
(http://creativecommons.org/licenses/by-nc-nd/4.0/).
J.-F. Côté, et al.
Nomenclature
asym crown asymmetry (m)
br branchiness index
cbh crown base height (m)
cpa crown projected area (m2)
cw crown width (m)
dbh diameter at breast height (cm)
dim fractal dimension
dist mean distance to its nearest neighbour (m)
fc fraction cover (%)
hcpa height to the maximum crown projected area (m)
hgt tree height (m)
hlc height to live crown (m)
knot knots surface area (m2)
la tree leaf area (m2)
lac lacunarity
lai leaf area index
nbr number of branches
nnr nearest neighbour ratio (m−1)
nwrl number of whorls
exposure, moisture and nutrient availability, stocking density, species
composition, and disturbance have direct impacts on wood properties
(Downes and Drew, 2008; Kint et al., 2010; Luss et al., 2019). At the
tree level, the individual growth rate, knot distribution and size, crown
geometry and branching structure vary significantly across a range of
species and sites (Mäkinen and Colin, 1998; Mansfield et al., 2007).
Tree size and architecture at a given age are strongly influenced by site
conditions and management. It is possible to predict tree structure at­
tributes from field inventory under certain conditions. For instance, in
pure Scot Pine stands branch characteristics can be predicted from tree-
level variables (such as stem diameter) without detailed knowledge of
the stand history (Mäkinen and Colin, 1998).
Characterization of wood fibre attributes (WFA) is important to
enhance the economic value of forest fibre, and once harvested, to di­
rect it to proper end-uses. This will ensure long-term sustainability of
the forest sector (MacKenzie and Bruemmer, 2009). For example, wood
density is a key attribute that influences strength and stiffness of lumber
products (Wegner et al., 2010; Hassegawa et al., 2019) as well as pulp
yield and the energy requirements for paper making processes. Micro­
fibril angle, fibre length, coarseness and modulus of elasticity are other
industrially important attributes that affect lumber stability and
strength and the strength and quality of paper products (Jozsa et al.,
1994). In forestry, understanding the development and variation of
wood quality is also important for several management and planning
problems (see Mäkelä et al., 2010 for a review). Wood quality models
provide information about the variation in wood properties that affect
the end-use of wood products. Industries need reliable assessment of
wood quality and its variation across forest sites at the time of harvest,
and the models must be capable of predicting wood properties directly
from measurable variables (e.g. Wilhelmsson et al., 2002; Ikonen et al.,
2008). Furthermore, the models of wood quality should apply to long
term silvicultural planning by linking with growth and yield model
outputs as well as being integrated into decision support systems (e.g.
Kantola et al., 2009; Newton, 2019).
Estimates of WFA rely on measurements of intrinsic or extrinsic
indicators of wood properties (van Leeuwen et al., 2011). Intrinsic in­
dicators relate to the internal, anatomical structure of the wood, the
proportions of juvenile and mature wood, as well as the cellular
structure. Although intrinsic indicators are crucial for mill operators,
they are rarely available over large forest estates before harvesting.
Typically, their measurements in radial and tangential planes require a
combination of microscopic digital imagery, X-ray densitometry, and X-
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Forest Ecology and Management 479 (2021) 118624
pai plant area index
pbf % of balsam fir
pbs % of black spruce
sd stem density (stems ha−1)
spc tree species
tap taper coefficient
vol tree volume (m3)
wal Weibull α-parameter for leaf area
waw Weibull α-parameter for wood volume
wbl Weibull β-parameter for leaf area
wbw Weibull β-parameter for wood volume
C coarseness (µg m−1)
FL fibre length (mm)
MFA microfibril angle (°)
MOE modulus of elasticity (Gpa)
RD radial diameter (µm)
SS specific surface (m2 kg−1)
TD tangential diameter (µm)
WD wood density (kg m−3)
WT wall thickness (µm)
ray diffraction (Evans, 2006; Downes et al., 2002; Downes and Drew,
2008). Extrinsic indicators of wood properties relate to the external
characteristics of a tree, including tree height, diameter, stem taper,
and crown structure. Recent studies explored the relationships between
intrinsic and extrinsic characteristics to assess wood fibre character­
istics from standing trees (Eriksson et al., 2006; Donaldson, 2008; Lenz
et al., 2012). Donaldson (2008) undertook detailed work relating WFA
to tree structure and form. They concluded that height and mechanical
stresses on the bole, such as wind and gravity, can have a direct impact
on WFA and thus serve as reliable indicators of wood properties.
The spatialization of WFA across forest landscapes makes exclusive
use of extrinsic indicators (Hilker et al., 2013). For instance, Lessard
et al. (2014) mapped a series of WFA at the landscape level across
Newfoundland, Canada, with climate and geographic data (e.g. tem­
perature, precipitation, elevation) and spatial layers from the provincial
forest inventory (e.g. stand age, species composition, height, site index).
The relationships obtained demonstrated the relevance of forest in­
ventory variables for estimating wood properties. Results also suggested
that improving the accuracy of forest structure and site description
would benefit WFA mapping. Concurrent research in developing en­
hanced forest inventories enable detailed spatial quantification of ter­
rain and vegetation at high spatial resolution across forest estates
(Woods et al., 2011; White et al., 2013; Luther et al., 2014; Furze et al.,
2017).
Remote sensing data and associated methods for estimating struc­
tural attributes of forests are continuously improving. For instance,
airborne lidar (a-lidar) systems can predict forest attributes at both plot-
and tree-level. The plot level prediction of forest attributes is based on a
statistical dependency between accurate measurements of forest height
and height variation derived from a-lidar data and response variables
measured from ground plots (Næsset and Økland, 2002; Kankare et al.,
2013; Bouvier et al., 2015). Tree-level inventories make use of a-lidar
data typically to detect individual treetops and to predict attributes of
interest using sets of allometric models. Algorithms and techniques si­
milar to those developed for aerial images can be used with a-lidar data
(Hyyppä et al., 2008). Tree-level inventories consist of a sequence of
steps that includes tree detection, feature extraction, and estimation of
tree attributes. An important consideration is that not all trees can
usually be detected. The degree to which individual trees are success­
fully detected depends on the detection algorithm and its para­
meterization (Kaartinen et al., 2012).
Terrestrial lidar (t-lidar) systems collect large amounts of 3D data on
J.-F. Côté, et al.
the fine-scale structure of trees and forest stands (Dassot et al., 2011).
For each t-lidar scan, millions of returned signals are registered, pro­
viding positional information on all elements surrounding the instru­
ment. Despite the potential for t-lidar to generate high quality data,
these data can be unreliable when acquired in natural forest environ­
ments. The main problem for a complete measurement of a specific
scene is the occlusion of the laser pulse. Groups of clustered or opaque
objects prevent detection of elements in occluded areas (Hopkinson
et al., 2004; Watt and Donoghue, 2005; Van der Zande et al., 2006; Van
der Zande et al., 2008). One way to reduce this problem is to merge
data from t-lidar scans taken from different points of view and perform
one-to-one scan alignment to produce a single co-registered point
cloud. However, this approach may create new problems of over­
sampling or misalignment. The issue of undersampling for objects lo­
cated farther from the instrument remains problematic. In addition, the
potential of using t-lidar to retrieve 3D structure is not fully exploited
due to other problems such as (i) movements generated by wind, (ii)
the presence of objects at a finer level than that which can be resolved
by the t-lidar, and (iii) misregistration of t-lidar positions. In addition to
these technical difficulties, t-lidar scans provide a raw sketch of the
spatial distribution of elements in 3D but do not give specific in­
formation on their geometry and connectivity (topology). Moreover, it
is not easy to distinguish between wood and foliage material from the
return signal with current instruments (Béland et al., 2014). Thus, even
when scans from different points of view are available, identification of
structural elements and construction of a topologically and geome­
trically correct 3D structure are far from trivial (Côté et al., 2011;
Raumonen et al., 2013; Hackenberg et al., 2014). Even if t-lidar data
appear rich in structural information, an architectural model may be
needed to overcome the limitations and produce a realistic 3D structure
from the point clouds.
Architectural models can provide a realistic description of the fine-
scale structure of forest stands. As an exact representation of the forest
structure is not possible in practice, 3D architectural models provide
convenient simplifications of forest structure at tree- or branch-
(Cescatti, 1997; Livny et al., 2011) and down to the individual conifer
shoot- or leaf-level (Xu et al., 2007; Pirk et al., 2012; Widlowski et al.,
2014). Representing tree structure is particularly challenging when
dealing with mature trees in complex environments (Runions et al.,
2007; Palubicki et al., 2009). In such cases, 3D architectural models
have to deal with a high level of competition for space, light and nu­
trient resources which increase the number of unknown variables re­
quired to reproduce realistic forest scenarios. The L-Architect (Lidar to
tree Architecture) modeling tool was developed to address the limita­
tions of t-lidar data in natural forest stands and to extract fine-scale
structural attributes of individual trees (Côté et al., 2009, 2011). L-
Architect uses geometrically registered t-lidar scans of individual trees
to construct a model of the tree structure. Expanding the model to
create a detailed representation of forest canopy structure requires the
creation of a catalog of individual tree samples scanned with a t-lidar
(Côté et al., 2012). L-Architect was compared to a series of field mea­
surements on the branching structure and foliage distribution of in­
dividual trees (Côté et al., 2013). It was further validated in a controlled
virtual environment, used to reconstruct forest plots, and applied to
create surrogate plots using tree attributes derived from a-lidar data
(Côté et al., 2018).
The fine-scale structural information possible with emerging lidar
technologies and methods (e.g. L-Architect) has potential to improve the
prediction and mapping of WFA. Following Lessard et al. (2014), two
studies refined the modeling of WFA in Newfoundland at the plot level
for balsam fir and black spruce forests with a-lidar (Luther et al., 2014)
and t-lidar (Blanchette et al., 2015). These studies improved upon the
level of spatial detail and previous relationships of Lessard et al. (2014),
whilst focusing on plot canopy structure, tree competition (with t-lidar)
and local topography. More recently, Giroud et al. (2019) predicted a
series of WFA at the individual tree level for black spruce in
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Forest Ecology and Management 479 (2021) 118624
Newfoundland by extracting tree metrics, crown characteristic and the
immediate tree’s surrounding from t-lidar.
Fernández et al. (2011) used an architectural model of radiata pine
to assess how wood quality in a timber species was linked to growth
processes during stand development. However, the study was limited to
young trees (up to 6 years old) and no independent measurements were
available to link the fibre attribute (wood density) with tree structure.
Lenz et al. (2012) demonstrated that morphological traits such as
height, crown dimensions and the distribution and size of live branches
at selected whorls explained almost 29% of the overall variation ob­
served in wood properties of white spruce (Picea glauca (Moench.)
Voss.). Pyörälä et al. (2019) inferred the variability of wood quality
indicators such as stem and crown descriptors from predicted tree
variables using terrestrial and airborne lidar for managed Scots pine
(Pinus sylvestris L.) stands in Finland. They identified the effect of ca­
nopy closure on crown longevity and stem growth, and the potential to
use stem and crown size variables predicted from lidar data to indicate
the development of size-dependent wood properties. However, it is still
unclear how better estimates of wood quality indicators can be
achieved using fine-scale structural information.
In this study, we used the validated architectural model L-Architect,
coupled with t-lidar data, to enhance the set of structural attributes
available for use as predictors of wood quality, beyond what is typically
available through forest inventory measurements. This raised the fol­
lowing research question: Does fine-scale structure of trees and forest
plots increase the variance of WFA that is explained with standard
forest inventory measurements?
Therefore, the main objective for this study was to evaluate the
extent to which we could improve prediction of WFA using fine-scale
structure derived from t-lidar data with L-Architect. First, we developed
predictive models for WFA at both tree- and plot-levels using para­
metric and non-parametric methods. Second, we assessed the impact of
fine-scale structure variables of trees and forest plots derived from L-
Architect on the predictive models. We hypothesized that fine-scale
structure would improve significantly the models’ predictions of WFA
(by at least 15% on average) when compared to predictions using only
in situ measurements.
2. Methods
2.1. In situ measurements
The study made use of a network of permanent sample plots (PSPs)
located in Newfoundland, Canada. The island of Newfoundland (ap­
proximately 111,000 km2) is situated at the eastern end of North
America’s Boreal Forest (Rowe, 1972). The forests of the island are
dominated by balsam fir (Abies balsamea (L.) Mill.), hereafter referred to
as fir, and black spruce (Picea mariana (Mill.) B.S.P.), hereafter referred
to as spruce with a limited variety of other species (Government of
Newfoundland and Labrador, 2019). The western part of the island has
considerable topography with peaks reaching up to 814 m from the
coastline. Pure stands of fir, which prefer moist and well-drained soils,
can attain heights of 20–24 m at 70–100 years on the best sites. How­
ever, growing conditions change significantly with altitude where sub-
arctic vegetation and dwarf patches of spruce and fir are found. Due to a
very high tolerance for unfavorable conditions, spruce is the dominant
species found over Central Newfoundland where the natural forest fire
regime contributes to its dominance. Hardwood stands do not represent
a major forest type over the island, however, trembling aspen (Populus
tremuloides Michx) and white birch (Betula papyrifera Marshall) stands
can be found on good growing sites such as in the river valleys of
western Newfoundland.
An extensive database was available to support the modeling of
WFA in Newfoundland. In this study, a subset of the larger dataset
collected and analyzed as part of the Newfoundland Fibre Inventory
Project was used (Lessard et al., 2014; Luther et al., 2014; Blanchette
J.-F. Côté, et al.
et al., 2015; Groot and Luther, 2015; Giroud et al., 2019). The WFA
were measured from wooden increment core samples taken at PSPs
maintained by the provincial forest management service of New­
foundland and Labrador (Newfoundland Forest Service, 2011). The
centre and four corner locations of each plot were measured using a
Trimble® GeoXHTM global positioning system (GPS). The GPS locations
were post-processed to provide plot locations with sub-metre accuracy.
At each plot, the species and diameter at breast height (dbh ), measured
at 1.3 m, were recorded for each tree above a specified size according to
plot maturity (Fig. 2, details in Supp., Table 1). Height (hgt ) was
measured for a sample of trees and species-specific relationships be­
tween dbh and hgt were developed according to Damman type (Meades
et al., 1994), and used to predict heights for all trees. Maps of tree
locations were created using, first, the precise GPS location of the PSP
central point. Then, the distance and azimuth of all trees of the plot
were measured from the PSP center using a Trimble® LaserAceTM 1000
rangefinder. The (x , y , z ) position of each measured tree was calculated
using the GPS location, distance, and azimuth information.
The wooden increment cores were extracted at breast height from
ten live trees selected outside the plots, at approximately ten meters
from each PSP at an angle of 135° from the corner where the site
conditions were the most representative of the PSP. Trees were selected
according to be representative of the range of dbh present within each
PSP (Fig. 2). Table 1 in the supplementary material details the char­
acteristics of the surveyed plots and their associated core sampled trees.
The wooden increment cores from the 10 trees per site were analyzed at
a state-of-the-art facility for wood and fibre analysis at the FPInnova­
tions laboratory in Vancouver (Goodison et al., 2007) using the Sil­
viScan system, which is a combination of optical microscopy, image
analysis, X-ray densitometry and X-ray diffractometry (Sherson et al.,
2007). WFA included wood density (WD ), radial and tangential fibre
diameter (RD , TD ), fibre coarseness (C ), microfibril angle (MFA ),
modulus of elasticity (MOE ), cell wall thickness (WT ) and specific fibre
surface (SS ). SS refers to the surface area per unit weight and as
coarseness, the SilviScan system calculates it using the cell dimensions
and WD estimates with a radial resolution of 25 µm. WT is determined
through WD and the rather constant tracheid wall density (Evans et al.,
1995). MOE for its part is determined with a semi-empirical approach
involving WD and the variation of the X-ray diffraction profile (Evans,
2006). In addition to the SilviScan data, WFA also comprised fibre
length (FL ). An estimate of each WFA for each site was calculated as a
weighted average of each wooden increment core measurement by
basal area of the sampled trees.
We made use of t-lidar data used in a coincident study on wood
quality modeling (Blanchette et al., 2015). A sample of 37 fir- and 31
spruce-dominated plots was selected to represent the range of species,
height, crown density and site quality classes present for the mer­
chantable forest area of the island, whereby the dominant species made
up more than 25% of the basal area of the plot (Fig. 1). Our t-lidar
sampling design consisted of scanning a subsample of the 680 available
core sampled trees located outside each plot. The selected trees scanned
with t-lidar were representative of the range of dbh and hgt of the
available core sampled trees (Fig. 2; Supp., Table 2). A total of 227
individual trees (82 spruce and 145 fir) from 68 plots were scanned
with the Ilris-3D (© Optech Inc.) in 2009 and the Imager 5006i (©
Zoller & Fröhlich, hereafter called Z + F) in 2010. We used the 227
trees scanned with t-lidar to reconstruct the structure of the trees and
the 68 PSPs using L-Architect as described in Section 2.2, as well as to
build tree level models of WFA. The Ilris-3D emits light at 1,500 nm and
scans with a maximum viewing window of 40° × 40°, which can be
adjusted to fit target objects. This system measures the position by
calculating time delay between the emission and the reception of signal
backscattering intensity in either first or last return mode. To reduce the
negative impact of signal occlusion, scans were taken from two to three
viewpoint locations around the tree using first-return modes. The trees
were scanned at a distance ranging from 10 to 15 m giving a point
4
Forest Ecology and Management 479 (2021) 118624
spacing1 (footprint2) of 5.0–7.5 mm (6.85–7.27 mm) respectively. The
Z + F emits light at 680 nm and scans with a near complete spherical
viewing window of 310° × 360°. The system uses the phase-shift
technology to measure distance for a maximum range of 80 m. We took
advantage of the very wide field of view to select the group of 10 live
merchantable trees to capture at the same time the trees and their
surrounding environments. Typically, five scans were performed within
and outside the tree group area to minimize occlusion: four scans lo­
cated outside (~North, South, East and West cardinal points) and one at
the center. The spatial resolution of the scans corresponded approxi­
mately to one point every 6.3 mm and a footprint3 of 4.1 mm on a
sphere at 10 m. The 3D point clouds taken by the Ilris-3D and Z + F
from different viewpoints were aligned into one geometric coordinate
system with the software Pointstream (3DImageSuite®) and Z + F La­
serControl respectively. Pointstream was then used to process all co-
registered point clouds to select and extract the individual trees used as
input for the architectural model L-Architect.
2.2. Tree and plot reconstruction
We used L-Architect to reconstruct Newfoundland fir and spruce
trees (Côté et al., 2013) and PSPs (Côté et al., 2018). L-Architect re­
produces realistic tree structures based on t-lidar data and allometric
relationships which define the total amount of foliage following two
main steps: branch growth and addition of foliage within the crown.
Specific rules for simulating the tree development are implemented at
the bud level. A two-step process construct models of realistic trees in
natural forest by determining space availability for branch growth and
by estimating light exposure for supporting foliage addition with a re­
source allocation model. For the tree reconstruction, we used specific
rules for tree development in conjunction with a space colonization
algorithm (Runions et al., 2007) to take into account the close en­
vironment of the branch. Foliage addition is based on shadow propa­
gation (Palubicki et al., 2009) that computes a coarse estimate of light
exposure for each bud to determine which ones produce new shoots.
The amount of light resources also serves to determine if a parent
branch supporting children is considered a liability for the tree and
should be shed. The branch diameters are estimated with a pipe model
proposed by Xu et al. (2007). For the plot reconstruction we used L-
Architect with in situ measurements of dbh and tree height (hgt ). We
chose for each tree recorded on a plot the best-fit co-registered scan that
matches the same species, site and crown density class, and minimizes
relative dbh and height differences between the scanned tree (subscript
‘s’) and the measured tree (subscript ‘m’), as follows:
|dbhm dbhs | |hgtm hgts |
+
dbhm hgtm (1)
Then, we scaled the selected tree scan in order to obtain the same
height of each tree found in the plot. The dbh , hgt and total leaf area (la )
estimated from allometric relationships (Lavigne et al., 1996; Newton,
2006) were inputs to L-Architect to create each tree model of a forest
plot. Finally, we made use of the tree map locations to position each
tree model in the plot.
2.3. Calculation of structural attributes
A series of structural attributes were selected to represent a wide
array of structures present at the tree and plot levels. We calculated
three types of structural attributes from the reconstructed plots: tree-
level attributes, plot-level attributes, and indicators of heterogeneity
1
Average angular resolution of 5 mm: arctan(6 mm/15000 mm) = 0. 23°
2
with RBeam is the mean radius (mm) and D is the distance to the target.
3
RBeam = (0.22 D + 6)/2 with RBeam is the mean radius (mm) and D is the
distance to the target.
J.-F. Côté, et al.
Fig. 1. Location of the permanent sample plots scanned with t-lidar in insular Newfoundland, Canada.
(Côté et al., 2018). Tree-level attributes included the total number of
branches (nbr ) and whorls (nwrl ), the maximum crown width (cw ), the
diameter at breast height (dbh ), the total height (hgt ), the height to live
crown (hlc ), the total knot surface (knot ) on the main stem, the total leaf
area (la), the branchiness ratio of the largest branch diameter to the dbh
(br ), the stem taper coefficient (tap ), the crown base height (cbh ), the
maximum crown projected area (cpa ), the height at cpa (hcpa ), the
crown volume (vol ) and the crown asymmetry (asym ). The plot-level
attributes included the fraction cover ( fc ), the leaf area index (lai ), the
plant area index ( pai ) as well as tree-level averaged attributes. We
characterized the vertical distributions of leaf area and wood volume by
fitting a two-parameters Weibull function:
x 1
W (x ; , ) = e ( / ) ,x 0,
(2)
where and are the scale and shape parameter. W (x ; , ) is zero for
x < 0 . The attributes wal , wbl , waw , wbw characterize the vertical dis­
tribution with the and parameter of leaf area and wood volume,
respectively. The stem density (sd ), the mean distance of a tree to its
nearest neighbour (dist ), and the nearest neighbour ratio (nnr ) char­
acterized the tree spatial competition at plot level:
5
Forest Ecology and Management 479 (2021) 118624
nnr = 2
dist
.
sd (3)
Fractal dimension and lacunarity quantified the heterogeneity of the
tree and plot structure. Fractal dimension dim is useful for studying
irregularity: it characterizes the way objects such as trees and forest
canopies occupy space. We used the box-counting method to estimate
fractal dimension of objects (Zeide and Pfeifer, 1991; Pradal et al.,
2009). We calculated the lacunarity index lac of the volume distribution
with a modified gliding box algorithm for 3D grey images (Plotnick
et al., 1996; Frazer et al., 2005; Kirkpatrick and Weishampel, 2005).
Lacunarity defines a scale-dependent deviation of a geometrical object
from translational invariance or homogeneity. Geometrical patterns
with low lacunarity are finely textured and homogeneous with respect
to the size distribution and spatial arrangement of gaps. In contrast,
patterns with high lacunarity show substantial heterogeneity in the size
distribution and spatial arrangement of gaps.
2.4. Model building and assessment
We used the fine-scale structural attributes calculated with L-
Architect to build predictive models of WFA at tree and plots levels.
J.-F. Côté, et al. Forest Ecology and Management 479 (2021) 118624

Fig. 2. Violin plots illustrating the distribution of forest attributes observed from plot and fibre tree measurements. Mean (point) and standard deviation (line) are
illustrated in red, while the median, interquartile range, percentiles (25th and 75th), minimum, maximum, and outliers are depicted in black by the overlaid
boxplots. Note: Species (%) is the proportion of plot basal area occupied by balsam fir or black spruce.

However, prior to building the predictive models, we selected a subset
of the available predictors based on several criteria. We discarded a
specific predictor if it had strong collinearity (|r| > 0.9) with one or
more predictor variables, had low correlation (|r| < 0.2 ) for all the WFA
response variables, or if there was strong evidence that the predictor
was not normally distributed through Shapiro–Wilk test (Shapiro and
Wilk, 1965): p-value < 0.01 and by visual inspection of the quantile-
quantile (Q-Q) plots.
We assessed the predictor importance in predicting WFA at tree and
plot levels with the algorithm Relieff (Robnik-Šikonja and Kononenko,
2003) and the function relieff implemented in the Statistics and
Machine Learning Toolbox of MATLAB (MathWorks v2019a). The Re­
lieff algorithm is a generic method for quantifying predictor im­
portance. The modified algorithm, RRelieff, can accommodate con­
tinuous predictors and can recognize nonlinear relationship between
predictors and the response variable. The algorithm penalizes the pre­
dictors that give different values to neighbors with the same response
values, and rewards predictors that give different values to neighbors
with different response values. The output of RRelieff algorithms is a set
of numerical weights representing the percentages of the predictors’
contribution to the variance of a response variable. Statistically, the
weight of a relevant predictor is expected to be positive and that of an
irrelevant one is expected to be zero or negative. We compared the
relative contribution of the fine-scale structure calculated with L-Ar­
chitect over the in situ measured inventory attributes to predict WFA.
We built four independent sets of predictive models at tree and plot
levels using two parametric and two non-parametric methods: namely
stepwise (SW) linear regression, best-subset (BS) linear regression,
random forest (RF) and partial least squares (PLS) regression. Each
method selects a subset of predictors according to different criteria.
SW regression uses a systematic method for adding and removing
terms from a linear model based on their statistical significance in ex­
plaining the response variable (Efroymson, 1960). The method begins
with an initial constant model, and then compares the explanatory

6
J.-F. Côté, et al.
power of incrementally larger (forward stepwise) and smaller (back­
ward stepwise) models to determine a final model. At each step, the
function searches for terms to add to or remove from the model based
on the Akaike information criterion (AIC) value (Akaike, 1973). The
function stepwiselm in the Statistics and Machine Learning Toolbox
of MATLAB implements the SW regression with several criteria options,
including AIC. We selected tree species (spc ) as a categorical variable
for the applicable tree-level models and kept the default values for the
other parameters.
BS regression consists of building a set of linear models where each
model contains a subset of the predictor variables (Hocking and Leslie,
1967; Miller, 1984; Heinze et al., 2018). Different implementations
exist based on all-subsets comparisons such as the “leap and bound”
algorithm (Furnival and Wilson, 1974). Here, we implemented a var­
iant where we build a total of 2P linear models, with P the number of
predictor variables. Let M0 denote the null model that contains no
predictors and simply predicts the sample mean of each observation.
()
P
Then for each q [0, P ], we built q models that contain exactly m
predictors. We selected a maximum of 10 best models among the q
P
()
models, called Mq , having the smallest leave-one-out cross validation
(LOOCV) error values. Then, we discarded models among the models
Mq indicating either heteroscedasticity, non-normality or autocorrela­
tion of their residuals and selected the final model based on the best R2
value. The function fitlm in the Statistics and Machine Learning
Toolbox of MATLAB implements linear regression models with a con­
stant term for the BS method. We selected spc as a categorical variable
for the applicable tree-level models and kept the default values for the
other parameters.
RF regression is an ensemble learning method that operates by
constructing a multitude of decision trees at training time and out­
putting the mean prediction of the individual trees (Breiman, 2001). A
random forest is a meta-estimator (i.e. it combines the result of multiple
Fig. 3. Correlogram of predictor (structure attributes) and response variables (wood fibre attributes) at tree level. The variable definitions are found in the no­
menclature.
7
Forest Ecology and Management 479 (2021) 118624
predictions) which aggregates many decision trees, with some mod­
ifications. The number of features split at each node is limited to some
percentage of the total number of features. This ensures that the en­
semble model does not rely too heavily on any individual feature, and
makes fair use of all potentially predictive features. Each tree draws a
random sample from the original data set when generating its splits,
adding a further element of randomness that prevents overfitting. In
this study, we used 500 trees whereby each tree was developed with a
random subset of the reference data and a random selection of pre­
dictors at each node of the tree to determine the split. The function
fitrensemble in the Statistics and Machine Learning Toolbox of MA­
TLAB implements a fit of ensemble of different learners for regression
including RF. The maximum number of decision splits was equal to the
number of observations minus one (N 1). The number of predictors to
select at random for each split was one third of the number of pre­
dictors. We selected spc as a categorical variable for the applicable tree-
level models and kept the default values for the other parameters.
PLS regression is a statistical method that bears some relation to
principal components regression. Instead of finding hyperplanes of
maximum variance between the response and independent variables, it
finds a linear regression model by projecting the predicted variables
and the observable variables to a new space (Mehmood et al., 2012). A
smaller number of latent components are identified that maximally
summarize the variation of the predictors while simultaneously re­
quiring maximum correlation with the response. PLS therefore strikes a
compromise between the objectives of reducing the predictor space
dimension and finding a predictive relationship with the response. PLS
regression is particularly suited when the matrix of predictors has more
variables than observations, and when there is multi-collinearity among
the predictor variables. By contrast, standard regression will fail in
these cases. We used the function plsregress in the Statistics and
Machine Learning Toolbox of MATLAB and chose the number of com­
ponents for each PLS models based on minimizing the mean square
J.-F. Côté, et al.
error of the predictions.
We assessed all four independent sets of predictive models using a
k -fold cross-validation due to the limited number of samples at both
tree and plot levels. We randomly partitioned the original sample into k
equal sized subsamples. Of the k subsamples, we kept one subsample for
testing the model, and used the remaining k 1 subsamples as training
data. The cross-validation process was then repeated k times, with each
of the k subsamples used exactly once as validation data. We used
k = 10 and summarized the cross-validation error statistics of both the
parametric and non-parametric models with the coefficient of de­
termination (R2 ), the normalized root mean squared error (NRMSE ) and
the normalized bias (NBias ), defined as:
N
(yi y¯) 2/ N
i=1
NRMSE = × 100
y¯
N NBias .
i=1
(yi y¯)/ N
NBias = × 100
y¯
For the parametric methods, we also evaluated the selected models
for heteroscedasticity (Breusch and Pagan, 1979), the non-normality
(Shapiro and Wilk, 1965) and the autocorrelation (Durbin and Watson,
1950) of the residuals.
2.5. Comparative analysis
We assessed the impact of fine-scale structure by comparing WFA
predictive models built with and without the fine-scale structure attri­
butes from L-Architect at tree and plot levels. We also assessed the im­
pact of including species information directly in the models or produ­
cing species-specific models to determine the extent to which the fine-
scale structure information captured the species information. At tree
level, the initial set of attributes consisted of 22 variables, with spc , dbh
and hgt measured in situ. At plot level, the initial set of attributes
consisted of 29 variables, including in situ measurement of tree-
Fig. 4. Correlogram of predictor (structure attributes) and response variables (wood fibre attributes) at plot level. The variable definitions are found in the no­
menclature.
Forest Ecology and Management 479 (2021) 118624
averaged values for dbh , hgt and stem density (sd ). We calculated the
plot species composition, summarized as the percentage of fir ( pbf ) and
spruce ( pbs ), from in situ measurements. Hereafter, we designate the
set of variables measured in situ, i.e. dbh , hgt and sd (at plot level), as
inventory attributes, and the set of fine-scale structure variables as L-
Architect attributes. Note that the inventory attributes are a subset of
the L-Architect attributes. The species-specific models consisted of only
balsam fir or black spruce trees at tree level, while the species-specific
models at plot level consisted of only plots dominated by the respective
species. We quantified the impact of L-Architect fine-scale structure on
the prediction of WFA with the triplet:
(i|j) = {R¯2i R¯2j ,NRMSE
¯ i ¯ i
¯ j , NBias
NRMSE ¯ j}
NBias (6)
that defines the mean (denoted as ‘bar’) difference between two sets of
(4) models, i and j , of the cross-validation error statistics R2 , NRMSE and
(5)
3. Results
3.1. Feature selection
Several predictor variables at both tree and plot levels failed the
Shapiro–Wilk test (Supp., Tables 2 and 3 show descriptive statistics of
predictor and response variables). Under a visual inspection of the Q-Q
plots, however, most of those variables did not show severe non-nor­
mality issue. Other variables exhibited strong collinearity with other
predictor variables or very little correlation with the response variables
(Figs. 3 and 4). After careful assessment, we decided to discard asym ,
cw , hlc , nbr , vol , wal , wbl at tree level, and cw , dist , lai , nbr , nwrl , pbf ,
vol , waw , wbw at plot level (Table 1). For consistency, we kept the same
set of predictors for all four statistical modeling methods even though
the non-parametric methods were less sensitive to collinearity and non-
normality issues.
We summarized in Figs. 5 and 6 the variable importance for the nine
8
J.-F. Côté, et al. Forest Ecology and Management 479 (2021) 118624

Table 1 regression. In the case of BS regressions, we were able to select models

Discarded predictor variables showing collinearity (|r| > 0.9) with other pre­ with no issues regarding the residuals. The parametric models were
dictor variables, correlation (|r| < 0.2) for all the WFA response variables, or stronger than the non-parametric models. For the non-parametric
non-normality (p-value < 0.01, and visual inspection of Q-Q plots). The methods, PLS regression models showed higher correspondence
variable definitions are found in the nomenclature.
(R2 [0.05, 0.80]) than those built with RF (R2 [0.01, 0.71]). Fig. 8
Discarded Predictor Rationale shows the scatter plots for the nine WFA attributes using PLS regression.
The observed versus predicted for the nine WFA attributes using PLS
Tree level
regression showed moderately strong, positive, linear correspondence
asym Non-normality (severe) and no correlation
cw Non-normality and collinearity with cpa with a few outliers, with the exception for FL that had weak corre­
hlc Non-normality (severe) and no correlation spondence. Based on PLS, we observed weak correspondence
nbr Non-normality and collinearity with nwrl (R2 < 0.24 ) for MFA and FL , moderate correspondence for , C , MOE
vol Non-normality and collinearity with cpa (R2 [0.36, 0.57]), while the other attributes, RD , WT , SS , and WD ,
wal Collinearity with waw
showed high correspondence (R2 > 0.66).
wbl Collinearity with wbw

Plot level
cw Collinearity with multiple predictors 3.4. Model comparison
dist Collinearity with nnr
lai Collinearity with pai
nbr Collinearity with multiple predictors The models developed using different combinations of predictors
nwrl Collinearity with multiple predictors served to quantify the impact of using fine-scale structure attributes
pbf Collinearity with pbs estimated with L-Architect in the models (Table 4). The results were
vol Non-normality and collinearity with cpa , knot consistent amongst the different regression techniques, but we reported
waw Collinearity with wal problems at selecting valid models for some WFA with parametric
wbw Collinearity with wbl
techniques, i.e. issues with non-normality, heteroscedasticity or auto­
correlation of the residuals. As a result, here we report the results based
WFA at tree and plot levels. At tree level, the six predictors with median on the models built with the PLS regressions.
weights greater than zero included information about leaf area (la ), At tree level, adding the fine-scale structure to the inventory attri­
stem form (tap ), branching structure (br ), material distribution (wbw , butes of dbh and hgt resulted in an improvement in the average error
waw ), and hgt (Fig. 5). Statistically, those were likely to contribute to a statistics of the 9 WFA (R2 , NRMSE and NBias ) with
(LA|I) = {0.23, 1.83, 0.03} . The best improvements were for SS , WD,
significant increase in performance for the predictive models. The dbh
and WT with differences in R2 > 40%, while improvement on MFA was
as well as the predictors related to spatial heterogeneity and crown
negligible.
geometry did not stand out as highly relevant features for most WFA.
Adding the tree species (spc ) as a categorical predictor variable
At plot level, the seven predictors with median weights greater than
improved the models using the inventory attributes by
zero included information about tree-level averaged la, crown geometry
(I + S|I) = {0.31, 2.67, 0.03} , and were closer to the models using the
attributes (hlc , cpa , asym ), plot-level attributes ( pai , fc , wbw ) and tree
L-Architect attributes, i.e. (LA|I + S) = { 0.08, 0.84, 0.00} . The gain of
positioning (nnr ) (Fig. 6). The other predictors, including inventory
adding spc to the models using L-Architect attributes was less significant
variables (dbh , hgt , sd ), as well as those related to material distribution,
with (LA + S|LA) = {0.08, 0.82, 0.01} . Species-specific, or stratified,
spatial heterogeneity and tree stem, had statistically less impact on the
tree level models did not improve with the L-Architect attributes with
performance of the predictive models for most WFA.
(LAfir |Ifir ) = {0.03, 0.29, 0.02} for balsam fir and
(LA spruce|Ispruce) = { 0.01, 0.05, 0.06} for black spruce.
3.2. Tree-level models

The tree-level models (Table 2) showed consistency across the four

statistical modeling methods R2 [0.13, 0.58], NRMSE [4.96, 29.12]
and NBias [ 0.04, 0.07]. For the parametric methods, we detected
several issues of non-normality, heteroscedasticity or autocorrelation of
the residuals. Within the set of models built with SW regression, three
failed at least two residual tests (MFA , MOE , WT ) and four failed one
(FL , RD , SS , TD , WD ). In the case of BS regressions, we were able to
select models having only minor issue with normality (MFA , RD , TD ,
WD ) or heteroscedasticity of the residuals (MFA ). For the non-para­
metric methods, the models built using PLS regression gave slightly
better error statistics for five of the nine attributes. The observed versus
predicted scatter plots for the nine WFA attributes using PLS regression
showed weak to moderately strong, positive, linear correspondence
with a few outliers, in particular for MFA , C , TD , and RD (Fig. 7). Based
on PLS, we observed weak correspondence (R2 < 0.24 ) for MFA , C , TD ,
and FL , while the other attributes, MOE , WT , SS , RD and WD , showed
moderate correspondence (R2 > 0.37 ).

3.3. Plot-level models
The plot-level models (Table 3) showed consistency across the four
statistical modeling methods R2 [0.10, 0.79], NRMSE [2.58, 15.68]
and NBias [ 0.08, 0.07]. There was only one parametric model that
failed the normality test on residuals, namely the MFA with SW
Fig. 5. Average predictor importance summarized for the nine WFA at tree
level using RRelieff. Variable importance expresses the contribution of pre­
dictors to explain WFA variance. On each box, the central mark indicates the
median, and the left and right edges of the box indicate the 25th and 75th
percentiles, respectively. The whiskers extend to the most extreme data points
not considered outliers, and the outliers are plotted individually using the '○'
symbol. The variable definitions are found in the nomenclature.

9
J.-F. Côté, et al. Forest Ecology and Management 479 (2021) 118624

including the L-Architect attributes did not improve by adding species

compositions, having (LA + SCfir |LAfir) = { 0.02, 0.11, 0.12} for balsam
fir and (LA + SCspruce |LA spruce) = {0.01, 0.01, 0.01} for black spruce
dominated plots.

4. Discussion

The estimation of internal wood quality such as fibre attributes is

Fig. 6. Average predictor importance summarized for the nine WFA at plot
level using RRelieff. Variable importance expresses the contribution of pre­
dictors to explain WFA variance. On each box, the central mark indicates the
median, and the left and right edges of the box indicate the 25th and 75th
percentiles, respectively. The whiskers extend to the most extreme data points
not considered outliers, and the outliers are plotted individually using the '○'
symbol. The variable definitions are found in the nomenclature.
At plot level, including the fine-scale structural improved the
average error statistics of the 9 WFA with (LA|I) = {0.33, 2.66, 0.04} .
The best improvements were for RD, SS , WD, and WT with differences
in R2 > 50%, while improvement on FL was negligible.
Adding the species composition variables (pbf and pbs ) improved
the inventory models by (I + SC|I) = {0.34, 2.70, 0.01} . These models
were comparable to models using L-Architect attributes, having
(LA|I + SC) = {0.01, 0.04, 0.05} . The gain of adding pbf and pbs to
the models using L-Architect attributes was negligible
(LA + SC|LA) = {0.01, 0.11, 0.01} . The species-specific plot level
models without species composition information benefited from the L-
Architect attributes with (LAfir |Ifir ) = {0.12, 0.21, 0.03} for balsam fir
and (LA spruce|Ispruce) = {0.13, 0.93, 0.10} for black spruce dominated
plots. Species-specific plot level models including species compositions
as predictor variables also improved with L-Architect attributes, with
(LA + SCfir |I+SCfir ) = {0.03, 0.13, 0.05} for balsam fir and
(LA + SCspruce |I+SCspruce) = {0.17, 1.10, 0.04} for black spruce domi­
nated plots. The results were very similar when compared to the stra­
tified models using L-Architect attributes, but without species compo­
sition, i.e. (LAfir |I+SC fir ) = {0.06, 0.24, 0.07} and
(LA spruce|I+SCspruce) = {0.16, 1.11, 0.05} for balsam fir and black
spruce dominated plots respectively. The stratifiedplot level models
challenging considering the numerous factors that determine the wood
fibre properties. Those factors include tree species, crown development,
stem shape, tree branchiness as well as stand conditions. Previous
studies examined details of the variability of wood quality as a function
of tree architecture (Fernández et al., 2011; Lenz et al., 2012; Pyörälä
et al., 2019). However, it was still unclear how better estimates of WFA
can be achieved using fine-scale structural information. The calibration
of L-Architect at both tree and plot level (Côté et al., 2013, 2018) proved
its ability to provide detailed structural information about individual
trees and coniferous forest plots composed of black spruce and balsam
fir. In this study, we not only demonstrated that fine-scale structural
information improves the accuracy of WFA model predictions, but that
the fine-scale attributes related to crown geometry, branching struc­
ture, stem forms, material distribution and spatial competition are
amongst the key factors in predicting WFA.
Finding suitable and valid models with parametric methods was
more problematic at tree-level, in particular through SW regression. We
tested several conditions to add or remove terms into the resulting
models; namely differences in p -value for F -test, changes in AIC or BIC
(Schwarz, 1978), and increases in R2 or adjusted R2 . We also limited the
model to linear and constant terms (no interactions). However, we
found only one model at tree-level without at least one severe issue
with the residuals tested over normality, heteroscedasticity and auto­
correlation. We implemented BS regression to allow a better control
over the model selection. We also tested different conditions during
model selection including LOOCV, AIC or BIC, Mallow’s Cp (Mallows,
1973) and R2 . We opted for a combination of minimizing LOOCV at
selecting the ten best models per class of q predictors and the best R2
satisfying (when possible) the three tests on residuals. As a con­
sequence, most of the tree level-models indicated only minor non-
normality issues using a strict Shapiro-Wilk test. These issues were not
present for plot level models. The non-parametric methods are not
sensitive to collinearity or to the number of predictors, and these issues
did not invalidate any of the tree or plot level models. PLS regression
gave on average similar or better results compared to RF using the
specified parameterization. In addition, the PLS built the 20 sets of
regression models in two minutes whilst RF (in parallel) took ap­
proximately two hours on a HP Workstation Z820 equipped with two
Intel® Xeon® CPU E5-2660 @ 2.20 GHz and 96 Gb DDR3-1866 ECC
RAM memory. This justified the use of PLS for comparing the sets of

Table 2
Predictive models for WFA at tree level. For stepwise and best subset regression, the following symbols indicate issues from residual assessments:
α = autocorrelation, η = heteroscedasticity, ν = normality. The out-of-bag loss regression error (OOBL) compares the out-of-bag predictions against the true
responses for all observations used for training. We chose the number of components for each partial least square models based on minimizing the mean square error
of the predictions. R2, NRMSE and NBias are cross-validated values. The WFA variable definitions are found in the nomenclature.
Stepwise Best subset Random forests Partial least square

R2 NRMSE NBias Issues R2 NRMSE NBias Issues R2 NRMSE NBias OOBL R2 NRMSE NBias Nb. Comp.

C 0.24 9.57 0.02 0.23 9.62 0.03 0.18 9.90 0.00 1310.20 0.18 10.01 0.00 5
FL 0.28 9.50 0.07 α 0.26 9.6 0.03 0.21 9.95 0.22 0.05 0.24 9.76 −0.02 2
MFA 0.15 28.58 0.05 ν,η,α 0.16 28.61 −0.04 ν,η 0.07 30.23 −0.06 19.37 0.12 29.06 0.15 2
MOE 0.43 17.46 0.03 η,α 0.44 17.36 −0.02 0.28 19.65 0.02 6.05 0.37 18.41 0.04 5
RD 0.55 5.61 0.00 ν 0.54 5.67 0.03 ν 0.56 5.56 0.06 2.59 0.52 5.79 −0.07 14
SS 0.49 9.11 0.03 α 0.46 9.39 0.06 0.46 9.43 −0.08 983.63 0.47 9.32 −0.08 5
TD 0.25 4.82 0.01 η 0.25 4.81 0.01 ν 0.17 5.13 −0.01 2.01 0.19 5.06 0.02 8
WD 0.58 10.48 0.02 ν 0.58 10.55 0.02 ν 0.58 10.48 −0.05 2499.50 0.56 10.81 0.04 5
WT 0.49 10.07 0.01 ν,α 0.47 10.28 0.03 0.47 10.31 −0.10 0.06 0.47 10.29 −0.10 7

10
J.-F. Côté, et al. Forest Ecology and Management 479 (2021) 118624

Fig. 7. Tree level observed vs. predicted values using a k -fold (k = 10 ) cross-validation for the nine WFA using partial least square regression.

models built with different sets of continuous and categorical pre­
dictors.
The models built with different predictor variables at tree or plot
level provided quantitative evidence of the impact of fine-scale struc­
ture attributes in model building. The best models used the structure
attributes estimated from the reconstructed trees or plots with archi­
tectural modeling. The general models improved by 23% and 33% on
average compared to those using inventory measurements without
species information at tree- and plot-levels respectively. At tree level,
the PLS regression models explained on average 20% and 11% of the
variation for balsam fir and black spruce species-specific WFA respec­
tively. Interestingly, species-specific models did not benefit from using
L-Architect attributes. This demonstrated that the L-Architect attributes
carry most of the species contribution to the WFA predictive tree level
models.
The structure measured from L-Architect reconstructed trees

Table 3
Predictive models for the WFA at plot level. For stepwise and best subset regression, the following symbols indicate issues upon residual assessments:
α = autocorrelation, η = heteroscedasticity, ν = normality. The out-of-bag loss regression error (OOBL) compares the out-of-bag predictions against the true
responses for all observations used for training. We chose the number of components for each partial least square models based on minimizing the mean square error
of the predictions. R2, NRMSE and NBias are cross-validated values. The WFA variable definitions are found in the nomenclature.
Stepwise Best subset Random forests Partial least square

R2 NRMSE NBias Issues R2 NRMSE NBias Issues R2 NRMSE NBias OOBL R2 NRMSE NBias Nb. Comp.

C 0.61 5.54 0.05 0.6 5.58 0.19 0.51 6.13 −0.23 499.18 0.50 6.26 −0.16 4
FL 0.14 6.68 −0.07 0.2 6.46 0.04 0.02 7.37 −0.04 0.03 0.05 7.02 −0.06 2
MFA 0.22 15.16 −0.17 ν 0.26 14.72 −0.15 0.01 17.68 −0.21 6.60 0.24 15.17 0.20 4
MOE 0.62 12.08 0.11 0.65 11.53 0.01 0.58 12.55 −0.40 2.44 0.57 12.79 −0.04 4
RD 0.76 2.88 0.08 0.76 2.92 0.1 0.53 4.04 −0.08 1.43 0.66 3.55 0.18 17
SS 0.80 5.51 −0.13 0.82 5.13 0.04 0.67 6.90 0.28 467.85 0.76 6.06 0.02 11
TD 0.44 2.47 −0.02 0.43 2.49 −0.06 0.33 2.71 −0.03 0.55 0.36 2.64 0.01 2
WD 0.83 6.10 0.08 0.82 6.32 0.04 0.69 8.13 −0.02 1565.50 0.80 6.80 0.08 19
WT 0.79 6.09 0.10 0.78 6.18 0.06 0.71 7.13 −0.34 0.03 0.73 6.85 −0.05 7

11
J.-F. Côté, et al.
Fig. 8. Plot level observed vs. predicted values using a k -fold (k = 10 ) cross-validation for the nine WFA using partial least square regression.
contributed significantly at discriminating species which in turn is one
of the key factors driving the tree wood fibre properties. Accordingly,
the set of non-stratified models using L-Architect attributes, including
tree species as a predictor, only improved the explained variation by
8%. At plot level, the PLS regression models explained 21% and 28% of
the WFA variation for balsam fir and black spruce dominated plots,
respectively. The inclusion of species composition did not improve the
results when using the L-Architect attributes. As for the tree level
models, the results demonstrated that the fine-scale structure carries a
significant part of the species composition contribution to the WFA
predictive plot level models.
The predictor importance assessment clearly demonstrated that the
most ranked variables were estimated from the architectural modeling
approaches. However, we observed that including species as predictors
in the regression models gave similar or slightly better results than
using L-Architect attributes alone. We explored the relationship between
structure and species predictors by using random forest to classify tree
and plot dominant species using fine-scale structure. The method is
found in the supplementary material. The topmost ranked predictors
(Supp., Fig. 1) were la and pai while other predictors estimated from
architectural modeling for tree (dbh, nwrl, hgt , wbw ) and dominant
(fc, cpa, wbl) species ranked in the top five. The results gave an overall
accuracy of approximately 88% either at classifying tree species or plot
dominant species (Supp., Fig. 2). This analysis demonstrated a strong
relationship between tree or forest canopy structure, and tree species or
12
Forest Ecology and Management 479 (2021) 118624
stand composition. The morphological differences between balsam fir
and black spruce explained this relationship, and we would expect to
find similar results for forests exhibiting comparable species composi­
tion conditions.
Adding L-Architect attributes for the non-stratified tree level models
explained 35% of the variation on average. The most challenging at­
tributes to predict were MFA , C , TD , and FL , compared to MOE , WT ,
SS , RD and WD . The difficulty of measuring accurately the former at­
tributes through wooden increment core analysis could explain in part
the lower explained variance. Interestingly, those former variables tend
to be under low genetic control in spruce compared with the latter
variables, which indicates a stronger environmental control (Lenz et al.,
2010). Nevertheless, we purposively discarded environmental and site
conditions that can have significant effects on WFA to focus solely on
the impact of tree structure. The results presented here are in agree­
ment with a previous study of black spruce in Newfoundland using
detailed t-lidar metrics of the tree, its crown, and its immediate en­
vironment, where the models explained 33%, 51%, 44%, 52% and 47%
of the variance in C , FL, MFA , MOE and WD (Giroud et al., 2019).
Here, the tree structure attributes were the only predictor variables
included in the models. Adding metrics relating to aspect of canopy
structure, competition, vegetation density, and local topography would
most likely improve the results, but would necessitate additional ana­
lysis that were not in the scope of this study.
The L-Architect attributes had a major impact on the non-stratified
J.-F. Côté, et al. Forest Ecology and Management 479 (2021) 118624

Table 4
Comparison of WFA predictive models developed using PLS regressions with different sets of predictor variables (LA: L-Architect, I: inventory, S: species, SC: species
composition, fir: fir or fir-dominated, spruce: spruce or spruce-dominated).
Model Set C FL MFA MOE RD

R2 NRMSE NBias R2 NRMSE NBias R2 NRMSE NBias R2 NRMSE NBias R2 NRMSE NBias

Tree-level analysis LA 0.18 10.01 0.00 0.24 9.76 −0.02 0.12 29.06 0.15 0.37 18.41 0.04 0.52 5.79 −0.07
LA + S 0.26 9.42 0.07 0.24 9.72 −0.04 0.12 29.19 0.11 0.46 16.95 −0.25 0.57 5.53 −0.08
LAfir 0.06 9.46 0.04 0.33 9.71 −0.08 0.17 30.80 0.02 0.29 16.47 −0.15 0.33 5.78 0.00
LAspruce 0.00 8.95 −0.02 0.03 9.55 0.01 0.10 24.51 0.51 0.18 16.89 −0.05 0.23 5.08 −0.02
I 0.02 10.83 0.02 0.20 9.97 −0.01 0.13 28.99 0.11 0.17 21.08 0.06 0.25 7.25 0.00
I+S 0.28 9.29 −0.05 0.23 9.78 0.03 0.12 29.08 0.04 0.44 17.26 0.07 0.58 5.43 −0.02
Ifir 0.03 9.59 0.03 0.33 9.71 −0.03 0.13 31.46 0.10 0.19 17.66 −0.07 0.36 5.65 −0.01
Ispruce 0.01 8.95 0.03 0.03 9.54 −0.03 0.10 24.29 0.07 0.16 17.03 −0.09 0.26 4.96 −0.02

Plot-level analysis LA 0.50 6.26 −0.16 0.05 7.02 −0.06 0.24 15.17 0.20 0.57 12.79 −0.04 0.66 3.55 0.18
LA + SC 0.50 6.29 −0.29 0.04 7.11 0.00 0.18 15.83 0.39 0.59 12.51 0.21 0.72 3.13 0.01
LAfir 0.06 5.98 −0.40 0.01 6.75 −0.08 0.09 15.35 −0.22 0.30 10.77 −0.32 0.28 2.85 0.02
LAspruce 0.46 5.45 −0.18 0.09 7.46 −0.90 0.44 13.77 0.51 0.52 10.19 −0.60 0.04 2.88 0.02
LA + SCfir 0.08 6.04 −0.18 0.00 6.97 −0.25 0.12 15.33 0.82 0.20 11.49 −0.08 0.27 2.89 0.03
LA + SCspruce 0.41 5.74 −0.76 0.02 7.64 −0.34 0.41 14.29 1.28 0.43 11.19 −1.06 0.00 2.80 −0.01
I 0.19 7.97 0.15 0.05 6.97 −0.10 0.19 15.45 −0.15 0.23 17.07 0.09 0.14 5.56 −0.08
I + SC 0.43 6.65 −0.05 0.02 7.22 −0.10 0.18 15.58 −0.25 0.59 12.39 0.11 0.78 2.75 0.02
Ifir 0.20 5.62 −0.01 0.04 6.65 −0.07 0.12 15.24 0.09 0.31 10.57 −0.19 0.08 3.20 0.03
Ispruce 0.14 6.91 0.00 0.14 6.95 −0.20 0.28 16.00 −0.23 0.32 12.37 −0.07 0.12 2.99 −0.17
I + SCfir 0.37 5.92 −0.06 0.00 6.98 −0.06 0.10 15.35 0.25 0.21 11.57 −0.22 0.18 3.10 −0.06
I + SCspruce 0.10 7.16 −0.29 0.04 7.43 −0.35 0.22 16.86 0.25 0.28 12.88 −0.54 0.04 2.66 0.06

Model Set SS TD WD WT

R2 NRMSE NBias R2 NRMSE NBias R2 NRMSE NBias R2 NRMSE NBias

Tree-level analysis LA 0.47 9.32 −0.08 0.19 5.06 0.02 0.56 10.81 0.04 0.47 10.29 −0.10
LA+S 0.61 8.01 −0.02 0.24 4.88 −0.01 0.71 8.70 −0.01 0.62 8.73 0.11
LAfir 0.10 8.11 −0.02 0.12 5.06 0.02 0.28 9.08 0.03 0.14 8.81 −0.01
LAspruce 0.10 7.33 0.02 0.03 4.54 0.03 0.23 7.62 0.00 0.08 8.18 0.01
I 0.05 12.48 −0.01 0.10 5.29 0.02 0.12 15.28 0.03 0.05 13.79 0.03
I+S 0.60 8.06 −0.01 0.23 4.88 0.01 0.72 8.57 −0.03 0.63 8.60 0.00
Lfir 0.07 8.25 −0.04 0.12 5.07 −0.02 0.23 9.40 0.05 0.08 9.06 0.03
Lspruce 0.09 7.34 0.02 0.04 4.51 −0.01 0.26 7.48 0.00 0.10 8.08 0.00

Plot-level analysis LA 0.76 6.06 0.02 0.36 2.64 0.01 0.80 6.80 0.08 0.73 6.85 −0.05
LA+SC 0.77 5.78 0.00 0.40 2.56 −0.01 0.82 6.34 −0.11 0.75 6.55 −0.14
LAfir 0.37 4.99 0.86 0.12 2.49 −0.05 0.28 5.72 −0.26 0.40 5.44 −0.16
LAspruce 0.30 6.52 0.67 0.00 2.37 −0.03 0.23 6.84 −0.16 0.40 6.42 −0.73
LA+SCfir 0.45 4.75 0.48 0.19 2.38 −0.02 0.33 5.56 −0.20 0.09 5.96 −0.07
LA+SCspruce 0.47 5.73 0.72 0.01 2.33 −0.05 0.33 6.31 −0.59 0.50 5.96 −0.52
I 0.22 10.71 −0.12 0.14 3.06 0.01 0.27 12.62 0.03 0.23 11.68 0.01
I+SC 0.74 6.14 0.10 0.51 2.31 −0.02 0.79 6.73 −0.12 0.72 7.01 0.04
Ifir 0.00 5.50 −0.04 0.01 2.63 −0.02 0.03 6.66 −0.04 0.00 6.20 −0.04
Ispruce 0.08 7.63 0.37 0.03 2.21 −0.06 0.08 7.39 −0.03 0.12 7.82 −0.07
I+SCfir 0.07 5.34 0.07 0.23 2.29 0.01 0.22 5.96 0.05 0.03 6.02 0.09
I+SCspruce 0.17 7.13 0.52 0.00 2.32 −0.09 0.06 7.64 −0.35 0.13 7.83 −0.16

plot models without species composition information, having 52% of
the variance explained on average. The reconstructed plots with L-
Architect thus allowed extracting attributes with strong relationships to
WFA, comparable to species composition (Lessard et al., 2014). Similar
to the tree level models, the MFA , FL, TD , and C attributes were also
those having the lowest correspondences. The same reasons mentioned
for tree level models may explain the difficulty to predict those attri­
butes. The strong predictions for attributes related to wood density and
mechanical wood stiffness (MOE , WT , SS , RD and WD ) at the tree and
plot level is encouraging for their inclusion into enhanced forest in­
ventories. Those traits are highly sought after by sawmills for the pro­
duction of lumber, which is the primary and most valuable transfor­
mation of conifer wood (Hassegawa et al., 2019). Our modelling results
are in agreement with the previous studies done in insular Newfound­
land, with R2 [0.35, 0.61] using environmental data and inventory for
C , FL, MFA , WD (Lessard et al., 2014), R2 [0.18, 0.53] using airborne
lidar for C , FL , MFA , MOE , RD , SS , WD , WT (Luther et al., 2014) and
R2 [0.37, 0.72] using terrestrial lidar for C , FL , MFA , WD (Blanchette
et al., 2015). Although, this study is unique by focusing on structure
attributes only. We deliberately discarded the site conditions, species
and environmental data from our predictor variables. It might be worth
further considerations to include attributes related to these aspects in
the future development of relationships between indirect remote sen­
sing data and wood quality.
Further modelling efforts might also consider a hierarchical ap­
proach to examine patterns of WFA. For example, Groot and Luther
(2015) explained portions of variance in wood density at ring, tree and
plot levels across Newfoundland with predictor variables at multiple
levels of a forest structural hierarchy (ring-, tree-, and plot-level vari­
ables) using linear mixed models (LMM). They concluded that there
was a strong influence of plot-level factors on wood density at all levels
of aggregation pointing to the need for enhanced inventory tools for
characterizing forest structure at plot and stand levels. We explored the
impact of grouping the structure attributes by spatial scale, i.e. tree and
plot level, using LMM. We present details and exploratory results in the
supplementary material. The LMM showed comparable results with
respect to the other modeling methods, and several predictors esti­
mated from L-Architect at both tree and plot levels were significant. The
analysis indicated that the plot identifier (random intercept) was sig­
nificant for all WFA. Regarding the hierarchical approach, Harrison

13
J.-F. Côté, et al.
et al. (2018) provide valuable recommendations regarding the difficulty
in selecting linear mixed models because of biases in the performance of
some tests (e.g. AIC comparisons) introduced by the presence of
random effects. Hence, a proper hierarchical analysis would require a
sampling design to ensure reliable quantification of the variance ex­
plained by the predictors, including their interactions, at each spatial
scale (tree, plot, landscape) for the set of WFA.
5. Conclusion
We investigated the contribution of architectural modeling for es­
timating WFA at the tree and plot levels for balsam fir and black spruce
trees growing in Newfoundland, Canada. This study demonstrated that
better characterization of the tree and plot structure using L-Architect
improved the predictive ability of the models for estimating WFA. We
assessed the impact of fine-scale structure on predictive models for WFA
with parametric and non-parametric methods. Non-parametric partial
least square regression provided the best compromise between robust­
ness on collinearity between predictors, predictive capability and
computation speed. A variable importance analysis demonstrated that
attributes related to crown geometry, branching structure, stem form,
canopy material distribution and spatial competition derived from L-
Architect were highly significant in the resulting models. The percentage
of variance explained, through cross-validation, ranged from 12–56%
and 5–80% at tree- and plot-levels respectively. The MFA , FL , TD , and
C were the most challenging attributes to predict at both levels. Due to
the difference in tree morphology, information on tree species and plot
composition did not make a significant difference in the predictive
capability for most of the WFA. The addition of fine-scale structure
improved the models built only with inventory attributes by 10–31%
and 0–53% at tree- and plot-levels respectively. These results provided
sufficient evidence to support our hypothesis with an average im­
provement of 23% and 33% at tree- and plot-levels respectively. Our
approach avoids the need for destructive measurements on tree struc­
ture while providing estimates at the same level of accuracy.
We demonstrated through a series of comparative analysis and
partitions over the set of predictors that L-Architect supports the de­
velopment of relationships between tree structure and its wood quality
characteristics. The results indicated that by using solely the fine-scale
characterisation of tree and plot structure with L-Architect, our results
compared with other available models developed with environmental
data, field measurements, and terrestrial and airborne lidar data. Our
method used very detailed structural dataset currently unavailable at
the stand level and provided enhanced capacity for forest inventory. We
demonstrated that a few key fine-scale structural attributes describing
the tree characteristics (asym , br , cpa , hlc , la , tap ), spatial competition
(nnr ) and canopy material distribution ( pai , waw , wbw ) would sig­
nificantly improve our predictive capacity for large-scale mapping of
wood quality, assuming that these attributes could be estimated using
above-canopy remote sensing data. The architectural modeling ap­
proach could further support, by integrating detailed structural data,
the development of empirical models and the retrieval of biophysical
properties with multi-scale remote sensing observations.
CRediT authorship contribution statement
Jean-François Côté: Conceptualization, Methodology, Software,
Validation, Formal analysis, Investigation, Resources, Writing - original
draft, Visualization, Supervision. Joan E. Luther: Conceptualization,
Writing - review & editing, Supervision, Resources, Project adminis­
tration, Funding acquisition. Patrick Lenz: Conceptualization,
Methodology, Writing - review & editing. Richard A. Fournier:
Writing - review & editing, Supervision, Resources, Funding acquisi­
tion. Olivier R. van Lier: Writing - review & editing, Resources, Data
curation, Visualization, Supervision.
14
Forest Ecology and Management 479 (2021) 118624
Declaration of Competing Interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influ­
ence the work reported in this paper.
Acknowledgments
The Newfoundland Fibre Project is a research initiative of the
Canadian Wood Fibre Centre (CWFC), Canadian Forest Service of
Natural Resources Canada, Corner Brook Pulp and Paper Limited
(CBPPL), FPInnovations, Grenfell Campus of Memorial University of
Newfoundland (MUN), NL Department of Natural Resources, Nova
Scotia Community College and the University of Sherbrooke. We thank
all participants of the project, notably, Dr. Wade W. Bowers of Grenfell
Campus (MUN). We thank Danny Blanchette and Emilie Lessard of the
University of Sherbrooke for their help during the data acquisition and
processing of the tree map locations and TLS scans. We also thank
Danny Blanchette for his help with the preliminary statistical analysis.
We thank François Rousseu of the University of Sherbrooke for his
advice on the use of several statistical approaches. We are also grateful
to Janet Bourque, Matthew Glover, Heidi Kavanagh, and Michael
Noonan of Grenfell Campus (MUN) for their help acquiring and pro­
cessing TLS scans. This work was supported by the Atlantic Canada
Opportunities Agency, Canada (ICF 786-16766-195492 to W.W.B.),
Centre for Forest Science and Innovation, Canada (221208 to W.W.B.),
Natural Science and Engineering Council of Canada (NSERC), Canada
(CRDPJ-390394 to R.A.F. and J.E.L., RGPIN-2014-04508 to R.A.F.),
Newfoundland and Labrador Department of Innovation, Business and
Rural Development, Canada (WES2010.06.02.045W to W.W.B.) and
Research and Development Corporation, Canada (5404.1221.102 to
W.W.B.). This research was also supported by the AWARE project
(NSERC CRDPJ-462973-14, grantee N.C. Coops, UBC), in collaboration
with the CWFC, FPInnovations and CBPPL.
Appendix A. Supplementary material
Supplementary data to this article can be found online at https://
doi.org/10.1016/j.foreco.2020.118624.
References
Akaike, H. (Ed.), 1973. Information theory and an extension of the maximum likelihood
principle. Akadémiai Kiadó, Budapest, Hungary.
Béland, M., Widlowski, J.L., Fournier, R.A., 2014. A model for deriving voxel-level tree
leaf area density estimates from ground-based LiDAR. Environ. Model. Software 51,
184–189.
Blanchette, D., Fournier, R.A., Luther, J.E., Côté, J.-F., 2015. Predicting wood fiber at­
tributes using local-scale metrics from terrestrial LiDAR data: A case study of
Newfoundland conifer species. For. Ecol. Manage. 347, 116–129.
Bouvier, M., Durrieu, S., Fournier, R.A., Renaud, J.-P., 2015. Generalizing predictive
models of forest inventory attributes using an area-based approach with airborne
LiDAR data. Remote Sens. Environ. 156, 322–334.
Breiman, L., 2001. Random forests. Mach. Learn. 45, 5–32.
Breusch, T.S., Pagan, A.R., 1979. A simple test for heteroscedasticity and random coef­
ficient variation. Econometrica 47, 1287–1294.
Cescatti, A., 1997. Modelling the radiative transfer in discontinuous canopies of asym­
metric crowns. II. Model testing and application in a Norway spruce stand. Ecol.
Model. 101, 275–284.
Côté, J.-F., Fournier, R.A., Egli, R., 2011. An architectural model of trees to estimate
forest structural attributes using terrestrial LiDAR. Environ. Model. Software 26,
761–777.
Côté, J.-F., Fournier, R.A., Frazer, G.W., Olaf Niemann, K., 2012. A fine-scale archi­
tectural model of trees to enhance LiDAR-derived measurements of forest canopy
structure. Agric. For. Meteorol. 166–167, 72–85.
Côté, J.-F., Fournier, R.A., Luther, J.E., 2013. Validation of L-Architect model for balsam
fir and black spruce trees with structural measurements. Can. J. Remote. Sens. 39,
S41–S59.
Côté, J.-F., Fournier, R.A., Luther, J.E., van Lier, O.R., 2018. Fine-scale three-dimensional
modeling of boreal forest plots to improve forest characterization with remote sen­
sing. Remote Sens. Environ. 219, 99–114.
Côté, J.-F., Widlowski, J.-L., Fournier, R.A., Verstraete, M.M., 2009. The structural and
J.-F. Côté, et al.
radiative consistency of three-dimensional tree reconstructions from terrestrial lidar.
Remote Sens. Environ. 113, 1067–1081.
Dassot, M., Constant, T., Fournier, M., 2011. The use of terrestrial LiDAR technology in
forest science: application fields, benefits and challenges. Ann. For. Sci. 68, 959–974.
Donaldson, L., 2008. Microfibril angle: measurement, variation and relationships – a
review. IAWA J. 29, 345–386.
Downes, G.M., Drew, D.M., 2008. Climate and growth influences on wood formation and
utilisation. South. For. 70, 155–167.
Downes, G.M., Wimmer, R., Evans, R., 2002. Understanding wood formation: gains to
commercial forestry through tree-ring research. Dendrochronologia 20, 37–51.
Durbin, J., Watson, G.S., 1950. Testing for serial correlation in least squares regression: I.
Biometrika 37, 409–428.
Efroymson, M., 1960. Multiple regression analysis. Math. Methods Digital Comput.
191–203.
Eriksson, D., Lindberg, H., Bergsten, U., 2006. Influence of silvicultural regime on wood
structure characteristics and mechanical properties of clear wood in Pinus sylvestris.
Silva Fenn. 40, 743–762.
Evans, R., Downes, G.M., Menz, D.N.J., Stringer, S.L., 1995. Rapid measurement of var­
iations in tracheid transverse dimensions in a radiata pine tree. Appita J. 48,
134–138.
Evans, R., 2006. Wood stiffness by X-ray diffractometry. In: Stokke, D., Groom, L. (Eds.),
Characterization of the cellulosic cell wall. Blackwell Publishing, Ames, Iowa, pp.
138–146.
Fernández, M.P., Norero, A., Vera, J.R., Pérez, E., 2011. A functional–structural model for
radiata pine (Pinus radiata) focusing on tree architecture and wood quality. Ann. Bot.
108, 1155–1178.
Frazer, G.W., Wulder, M.A., Niemann, K.O., 2005. Simulation and quantification of the
fine-scale spatial pattern and heterogeneity of forest canopy structure: A lacunarity-
based method designed for analysis of continuous canopy heights. For. Ecol. Manage.
214, 65–90.
Furnival, G.M., Wilson, R.W., 1974. Regressions by Leaps and Bounds. Technometrics 16,
499–511.
Furze, S., Ogilvie, J., Arp, P.A., 2017. Fusing digital elevation models to improve hy­
drological interpretations. J. Geogr. Inf. Syst. 9, 558–575.
Giroud, G., Schneider, R., Fournier, R.A., Luther, J.E., Martin-Ducup, O., 2019. Modeling
black spruce wood fiber attributes with terrestrial laser scanning. Can. J. For. Res. 49,
661–669.
Goodison, A., Sherson, G., Huntley, S., 2007. EvaluTree - A new resource for the pulp and
paper industry in Canada. Pulp Pap. Canada 108, 18–20.
Government of Newfoundland and Labrador, 2019. Forest Types | Forestry and Agrifoods
Agency. Available online: https://www.faa.gov.nl.ca/forestry/our_forest/forest_
types.html (accessed on May 4th 2020).
Groot, A., Luther, J.E., 2015. Hierarchical analysis of black spruce and balsam fir wood
density in Newfoundland. Can. J. For. Res. 45, 805–816.
Hackenberg, J., Morhart, C., Sheppard, J., Spiecker, H., Disney, M., 2014. Highly accurate
tree models derived from terrestrial laser scan data: a method description. Forests 5,
1069–1105.
Harrison, X.A., Donaldson, L., Correa-Cano, M.E., Evans, J., Fisher, D.N., Goodwin,
C.E.D., Robinson, B.S., Hodgson, D.J., Inger, R., 2018. A brief introduction to mixed
effects modelling and multi-model inference in ecology. PeerJ 6 (e4794), 1–32.
Hassegawa, M., Savard, M., Lenz, P.R.N., Duchateau, E., Gélinas, N., Bousquet, J., Achim,
A., 2019. White spruce wood quality for lumber products: priority traits and their
enhancement through tree improvement. Forestry 93, 16–37.
Heinze, G., Wallisch, C., Dunkler, D., 2018. Variable selection - A review and re­
commendations for the practicing statistician. Biom J 60, 431–449.
Hilker, T., Frazer, G.W., Coops, N.C., Wulder, M.A., Newnham, G.J., Stewart, J.D., van
Leeuwen, M., Culvenor, D.S., 2013. Prediction of wood fiber attributes from LiDAR-
derived forest canopy indicators. For. Sci. 59, 231–242.
Hocking, R.R., Leslie, R.N., 1967. Selection of the best subset in regression analysis.
Technometrics 9, 531–540.
Hopkinson, C., Chasmer, L., Young-Pow, C., Treitz, P., 2004. Assessing forest metrics with
a ground-based scanning lidar. Can. J. For. Res. 34, 573–583.
Hyyppä, J., Hyyppä, H., Leckie, D., Gougeon, F., Yu, X., Maltamo, M., 2008. Review of
methods of small-footprint airborne laser scanning for extracting forest inventory
data in boreal forests. Int. J. Remote Sens. 29, 1339–1366.
Ikonen, V.-P., Peltola, H., Wilhelmsson, L., Kilpeläinen, A., Väisänen, H., Nuutinen, T.,
Kellomäki, S., 2008. Modelling the distribution of wood properties along the stems of
Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) as affected
by silvicultural management. For. Ecol. Manage. 256, 1356–1371.
Jozsa, L.A., Middleton, G.R., Corp, F.C., 1994. A discussion of wood quality attributes and
their practical implications. Forintek Canada Corp., Western Laboratory.
Kaartinen, H., Hyyppä, J., Yu, X., Vastaranta, M., Hyyppä, H., Kukko, A., Holopainen, M.,
Heipke, C., Hirschmugl, M., Morsdorf, F., Næsset, E., Pitkänen, J., Popescu, S.,
Solberg, S., Wolf, B.M., Wu, J.-C., 2012. An international comparison of individual
tree detection and extraction using airborne laser scanning. Remote Sens. 4, 950–974.
Kankare, V., Vastaranta, M., Holopainen, M., Räty, M., Yu, X., Hyyppä, J., Hyyppä, H.,
Alho, P., Viitala, R., 2013. Retrieval of forest aboveground biomass and stem volume
with airborne scanning LiDAR. Remote Sens. 5, 2257–2274.
Kantola, A., Song, T., Usenius, A., Heikkilä, A., 2009. Simulated yield and quality dis­
tribution of sawn timber from final felling in a Norway spruce [Picea abies (L.) Karst.]
stand with varying thinning regimes: A case study. Wood. Mater. Sci. Eng. 4, 87–97.
Kint, V., Hein, S., Campioli, M., Muys, B., 2010. Modelling self-pruning and branch at­
tributes for young Quercus robur L. and Fagus sylvatica L. trees. For. Ecol. Manage.
260, 2023–2034.
Kirkpatrick, L.A., Weishampel, J.F., 2005. Quantifying spatial structure of volumetric
neutral models. Ecol. Model. 186, 312–325.
15
Forest Ecology and Management 479 (2021) 118624
Lavigne, M.B., Luther, J.E., Franklin, S.E., Jr, E.R.H., 1996. Comparing branch biomass
prediction equations for Abies balsamea. Can. J. For. Res. 26, 611–616.
Lenz, P., Bernier-Cardou, M., MacKay, J., Beaulieu, J., 2012. Can wood properties be
predicted from the morphological traits of a tree? A canonical correlation study of
plantation-grown white spruce. Can. J. For. Res. 42, 1518–1529.
Lenz, P., Cloutier, A., MacKay, J., Beaulieu, J., 2010. Genetic control of wood properties
in Picea glauca — an analysis of trends with cambial age. Can. J. For. Res. 40,
703–715.
Lessard, E., Fournier, R.A., Luther, J.E., Mazerolle, M.J., van Lier, O.R., 2014. Modeling
wood fiber attributes using forest inventory and environmental data for
Newfoundland’s boreal forest. For. Ecol. Manage. 313, 307–318.
Livny, Y., Pirk, S., Cheng, Z., Yan, F., Deussen, O., Cohen-or, D., Chen, B., 2011. Texture-
lobes for tree modelling. ACM Trans. Graph. 30 (53), 1–10.
Luss, S., Lundqvist, S.-O., Evans, R., Grahn, T., Olsson, L., Petit, G., Rosner, S., 2019.
Within-ring variability of wood structure and its relationship to drought sensitivity in
Norway spruce trunks. IAWA J. 40, 288–310.
Luther, J.E., Skinner, R., Fournier, R.a., Van Lier, O.R., Bowers, W.W., Coté, J.F.,
Hopkinson, C., Moulton, T., 2014. Predicting wood quantity and quality attributes of
balsam fir and black spruce using airborne laser scanner data. Forestry 87, 313–326.
MacKenzie, J., Bruemmer, G., 2009. Enhancing Canada's forest fibre. For. Chron. 85,
353–354.
Mäkelä, A., Grace, J., Deckmyn, G., Kantola, A., Kint, V., 2010. Simulating wood quality
in forest management models. For. Syst. 19, 48–68.
Mäkinen, H., Colin, F., 1998. Predicting branch angle and branch diameter of Scots pine
from usual tree measurements and stand structural information. Can. J. For. Res. 28,
1686–1696.
Mallows, C.L., 1973. Some comments on CP. Technometrics 15, 661–675.
Mansfield, S.D., Parish, R., Goudie, J.W., Kang, K.-Y., Ott, P., 2007. The effects of crown
ratio on the transition from juvenile to mature wood production in lodgepole pine in
western Canada. Can. J. For. Res. 37, 1450–1459.
Meades, W.J., Moores, L., Canada-Newfoundland Forest Resource Development, A.,
Canada, Forestry, C., Newfoundland, Labrador, R., Department of, F., Agriculture,
1994. Forest site classification manual: a field guide to the Damman forest types of
Newfoundland. Western Newfoundland Model Forest, Inc., Corner Brook, Nfld.
Mehmood, T., Liland, K.H., Snipen, L., Sæbø, S., 2012. A review of variable selection
methods in Partial Least Squares Regression. Chemometrics Intellig. Lab. Syst. 118,
62–69.
Miller, A.J., 1984. Selection of Subsets of Regression Variables. J. R. Stat. Soc. Ser. A. Stat.
Soc. 147, 389–425.
Næsset, E., Økland, T., 2002. Estimating tree height and tree crown properties using
airborne scanning laser in a boreal nature reserve. Remote Sens. Environ. 79,
105–115.
Newfoundland Forest Service, 2011. Permanent sample plot program procedures and
specifications. 2011 Revision. Newfoundland Forest Service, Forest Ecosystem
Management Division, Corner Brook, NL.
Newton, P.F., 2006. Intraspecific competition processes and their management within
black spruce (Picea mariana (Mill.) B.S.P.) stands. Faculty of Forestry. University of
British Columbia, Vancouver, British Columbia, Canada.
Newton, P.F., 2019. Wood quality attribute models and their utility when integrated into
density management decision-support systems for boreal conifers. For. Ecol. Manage.
438, 267–284.
Palubicki, W., Horel, K., Longay, S., Runions, A., Lane, B., Měch, R., Prusinkiewicz, P.,
2009. Self-organizing tree models for image synthesis. ACM Trans. Graph. 28 (58),
1–10.
Pirk, S., Stava, O., Kratt, J., Said, M.A.M., Neubert, B., Měch, R., Benes, B., Deussen, O.,
2012. Plastic Trees: Interactive Self-Adapting Botanical Tree Models. ACM Trans.
Graph. 31 (50), 1–10.
Plotnick, R., Gardner, R., Hargrove, W., Prestegaard, K., Perlmutter, M., 1996. Lacunarity
analysis: A general technique for the analysis of spatial patterns. Phys. Rev. E 53,
5461–5468.
Pradal, C., Boudon, F., Nouguier, C., Chopard, J., Godin, C., 2009. PlantGL: A Python-
based geometric library for 3D plant modelling at different scales. Graph. Models 71,
1–21.
Pyörälä, J., Saarinen, N., Kankare, V., Coops, N.C., Liang, X., Wang, Y., Holopainen, M.,
Hyyppä, J., Vastaranta, M., 2019. Variability of wood properties using airborne and
terrestrial laser scanning. Remote Sens. Environ. 235 (111474), 1–14.
Raumonen, P., Kaasalainen, M., Åkerblom, M., Kaasalainen, S., Kaartinen, H., Vastaranta,
M., Holopainen, M., Disney, M., Lewis, P., 2013. Fast automatic precision tree models
from terrestrial laser scanner data. Remote Sens. 5, 491–520.
Robnik-Šikonja, M., Kononenko, I., 2003. Theoretical and empirical analysis of ReliefF
and RReliefF. Mach. Learn. 53, 23–69.
Rowe, J.S., 1972. Forest Regions of Canada. Natural Resources Canada. Canadian Forest
Service, Publ. No. 1300.
Runions, A., Lane, B., Prusinkiewicz, P., 2007. Modeling trees with a space colonization
algorithm. In: Proceedings of the Third Eurographics conference on Natural
Phenomena. Eurographics Association, Prague, Czech Republic, pp. 63–70.
Schwarz, G., 1978. Estimating the dimension of a model. Ann. Statist. 6, 461–464.
Shapiro, S.S., Wilk, M.B., 1965. An analysis of variance test for normality (complete
samples). Biometrika 52, 591–611.
Sherson, G., Woo, K., Jang, H., Huntley, S., Drummond, J., da Silva Júnior, F., 2007. From
forest to product: new solutions for rapid,comprehensive wood and fibre analyses. In,
III ICEP – International Colloquium on Eucalyptus Pulp, p. 1–12.
Smith, W.K., Hinckley, T.M., 1995. Resource physiology of conifers: acquisition, alloca­
tion, and utilization. Academic Press, San Diego.
Väisänen, H., Kellomäki, S., Oker-Blom, P., Valtonen, E., 1989. Structural development of
Pinus sylvestrís stands with varying initial density: A preliminary model for quality of
J.-F. Côté, et al.
sawn timber as affected by silvicultural measures. Scand. J. For. Res. 4, 223–238.
Van der Zande, D., Hoet, W., Jonckheere, I., van Aardt, J., Coppin, P., 2006. Influence of
measurement set-up of ground-based LiDAR for derivation of tree structure. Agric.
For. Meteorol. 141, 147–160.
Van der Zande, D., Jonckheere, I., Stuckens, J., Verstraeten, W.W., Coppin, P., 2008.
Sampling design of ground-based lidar measurements of forest canopy structure and
its effect on shadowing. Can. J. For. Res. 34, 526–538.
van Leeuwen, M., Hilker, T., Coops, N.C., Frazer, G., Wulder, M.A., Newnham, G.J.,
Culvenor, D.S., 2011. Assessment of standing wood and fiber quality using ground
and airborne laser scanning: A review. For. Ecol. Manage. 261, 1467–1478.
Watt, P.J., Donoghue, D.N.M., 2005. Measuring forest structure with terrestrial laser
scanning. Int. J. Remote Sens. 26, 1437–1446.
Wegner, T., Skog, K.E., Ince, P.J., Michler, C.J., 2010. Uses and Desirable Properties of
Wood in the 21st Century. J. For. 108, 165–173.
White, J., Wulder, M., Vastaranta, M., Coops, N., Pitt, D., Woods, M., 2013. The utility of
16
Forest Ecology and Management 479 (2021) 118624
image-based point clouds for forest inventory: a comparison with airborne laser
scanning. Forests 4, 518–536.
Widlowski, J.-L., Côté, J.-F., Béland, M., 2014. Abstract tree crowns in 3D radiative
transfer models: Impact on simulated open-canopy reflectances. Remote Sens.
Environ. 142, 155–175.
Wilhelmsson, L., Arlinger, J., Spångberg, K., Lundqvist, S.-O., Grahn, T., Hedenberg, Ö.,
Olsson, L., 2002. Models for Predicting Wood Properties in Stems of Picea abies and
Pinus sylvestris in Sweden. Scand. J. For. Res. 17, 330–350.
Woods, M., Pitt, D., Penner, M., Lim, K., Nesbitt, D., Etheridge, D., Treitz, P., 2011.
Operational implementation of a LiDAR inventory in Boreal Ontario. For. Chron. 87,
512–528.
Xu, H., Gossett, N., Chen, B., 2007. Knowledge and heuristic-based modeling of laser-
scanned trees. ACM Trans. Graph. 26 (19), 1–13.
Zeide, B., Pfeifer, P., 1991. A method for estimation of fractal dimension of tree crowns.
For. Sci. 37, 1253–1265.