Southern Forests: a Journal of Forest Science

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Density effect on growth variables of eucalypt

clones in a Nelder wheel experiment

Franciele Alba da Silva, Sylvio Péllico Netto, Alexandre Behling, Henrique

Soares Koehler, Ataídes Marinheski Filho & Cláudio Cerqueira

To cite this article: Franciele Alba da Silva, Sylvio Péllico Netto, Alexandre Behling, Henrique
Soares Koehler, Ataídes Marinheski Filho & Cláudio Cerqueira (2022): Density effect on growth
variables of eucalypt clones in a Nelder wheel experiment, Southern Forests: a Journal of Forest
Science, DOI: 10.2989/20702620.2022.2040925

To link to this article: https://doi.org/10.2989/20702620.2022.2040925

Published online: 25 May 2022.

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Southern Forests 2022, 84(2): 01–12 Copyright © NISC (Pty) Ltd
Printed in South Africa — All rights reserved SOUTHERN FORESTS
This is the final version of the article that is ISSN 2070-2620 EISSN 2070-2639
published ahead of the print and online issue https://doi.org/10.2989/20702620.2022.2040925

Density effect on growth variables of eucalypt clones in a Nelder wheel

experiment
Franciele Alba da Silva 1 *  , Sylvio Péllico Netto 1  , Alexandre Behling 1  , Henrique Soares Koehler 2  ,
Ataídes Marinheski Filho 3  and Cláudio Cerqueira 3, 4 

1 Federal University of Paraná, Department of Forest Science, Curitiba, Paraná, Brazil

2 Federal University of Paraná, Department of Phytotechnology and Plant Health, Paraná, Brazil
3 Brasil Florestal, Canoinhas, Santa Catarina, Brazil

4 Eco Brasil Florestas, Araguaína, Tocantins, Brazil

* Corresponding author: francielealba47@gmail.com

Nelder’s systematic design enables studies of tree plantation spacing in relatively small areas, which is
of great interest for forest planning. However, dependence between observations can arise with this method,
making it difficult to use classic statistics for evaluating an experiment. Therefore, our aims were to: evaluate
the relationship between dendrometric variables (tree diameter, height and volume) and the available growing
area; evaluate autocorrelation among the observations; and use multivariate analysis of the profiles of the
clones for comparisons of the dendrometric variables. The data are from an experiment with four Nelder wheel
plots of different Eucalyptus species clones of different ages, in the state of Tocantins, Brazil. Spearman’s rank
correlation was applied to evaluate the relationship between the estimates of the studied variables and respective
available growing area. Autocorrelation between the observations was evaluated with Moran’s I index. To analyse
the profile behaviour of the clones, three questions were proposed: (1) Are there statistical differences between
the estimations made for the different clones? (2) Are there differences between treatments applied in the
experiment with the clones? (3) Is there interaction between clones and spacings? The main results show that
multivariate profile analysis solved the autocorrelation problem in Nelder’s systematic design. Furthermore, we
obtained answers as to differences between the growth variables diameter, volume and height among the clones
at different spacings, whether there is a difference between the spacings for these variables, and whether there is
an interaction between clones and spacings. The analysis method used for repeated measures is appropriate to
evaluate trees in Nelder wheel trials.

Keywords: autocorrelation, dendrometric variables, Eucalyptus platyphylla, Eucalyptus urograndis, experimental planting, multivariate profile
analysis, plantation spacing

Introduction

Tree spacing is an important factor in plantations because
it exerts an influence on the productivity of the trees
and the quality of the wood (Pretzsch and Rais 2016).
Experimental planting is useful for choosing tree spacing
(Stape and Binkley 2010). When new areas are included
for establishment of forest stands or new genotypes are
introduced, it opens space for studies that identify the main
effects of spacing on the growth and productivity of the
selected species (Gonçalves et al. 2008). The systematic
experimental design proposed by Nelder (1962) makes it
possible to test a high number of spacings because of the
gradual increase of distances between trees from a certain
point on, in relatively small areas (Parrott et al. 2012).
Due to its good characteristics, the design is attractive
for these studies and has bolstered research on many
forest species (Kuehne et al. 2013; Vanclay et al. 2013;
Uhl et al. 2015; Pachas et al. 2018) as well as eucalypt
species (Mabvurira and Miina 2002; Stape and Binkley
2010; du Plessis and Kotze 2011). However, the lack
of randomisation in Nelder’s systematic design makes
it difficult to use conventional analyses to evaluate it
statistically (Parrott et al. 2012). The independence of the
data is not guaranteed owing to a lack of randomisation of
the experiment, whereas the arrangement of the trees in
the spacing on an ordered scale increases the possibility
of autocorrelation. In Nelder’s designs, autocorrelation
can exhibit patterns of similarities or differences in sizes,
wherein trees at wider spacings exhibit special positive
correlation (Kim et al. 2015), while at closer spacings
and in regions with greater spacing heterogeneity the
autocorrelation is negative (Bullock and Burkhart 2005;
Fortin 2005; Kim et al 2015), with a contrasting size
hierarchy in their growth (Ford 1975).
These characteristics must be considered in evaluations
of Nelder wheel trials because ignoring the autocorrelation
aspect violates basic hypotheses of classical statistical
methods and can lead to biased results (Uhl et al. 2015);
therefore, it becomes important to use other statistical
techniques to carry out accurate analyses of the
experiments in a Nelder wheel plot. Many methods have

Southern Forests is co-published by NISC (Pty) Ltd and Informa UK Limited (trading as Taylor & Francis Group)
2 Alba da Silva, Netto, Behling, Koehler, Filho and Cerqueira
been used to control the autocorrelation problem in this
type of experiment. Aphalo and Rikala (2006) applied
generalised least-squares estimation with the error
variance modelled as a function of a covariate logarithm
of plant density. Affleck (2001) suggested alternative
analysis methods based on correlated spatial error
models to be used. Uhl et al. (2015), to better capture the
outcome of these interactions on tree growth in a Nelder
wheel considering spacing, employed general additive
regression models. Parrot et al. (2012) indicated the use
of mixed-effects models to circumvent the problem of
autocorrelation in Nelder wheels.
An alternative to solve the autocorrelation problem in
Nelder wheels is multivariate profile analysis. This analysis
is the multivariate equivalent of repeated measures or
mixed ANOVA and evaluates data that show autocorrelation
because of sequential measurements in space or over a
given period (Rencher 2002). Profile analysis provides an
alternative approach to analyse repeated measures taken on
the same experimental unit (Liu 2016).
In forestry, multivariate profile analysis has been used
to analyse the differentiation of annual height increments
conditioned on tree age (Zawieja and Kaźmierczak 2014),
in simultaneous statistical analysis of height curves
(Lappi 1997), and to investigate relative differences in
the growth of Sitka spruce and Norway spruce seedlings
under ozone exposure during three growing seasons
(Lukas and Diggle 1997).
Data analysis of Nelder wheel experiments using
multivariate profile analysis is appropriate to compare the
same dependent variables between groups in the space
points, which are autocorrelated. Furthermore, it is a simple
analysis applied to clearly express the results of interactions
between dendrometric variables, so as to compare clones
at different spacings. For example, to analyse dendrometric
data for J clones at K spacings in a Nelder wheel plot,
multivariate profile analysis is applied to evaluate three null
hypotheses (Cengiz 2020), as follows:
1. Parallelism: There is no interaction effect between
the J clones with the K spacings (segments) on the
dendrometric variables diameter, height and volume.
In this hypothesis, it is stated that the behaviour of the
means for the J clones is the same in the K spacings.
Graphically, the means for the J clones are parallel,
meaning that the profiles are parallel if the slopes for
each segment are the same (Rencher 2002).
2. Coincidence: There is no difference between the J
clones for the dendrometric variables diameter, height
and volume. In this hypothesis, it is stated that the
means of the dendrometric variables between the
clones do not differ from each other.
3. Flat: There is no difference between the K spacings
for the dendrometric variables diameter, height and
volume. This null hypothesis is evaluated for each J
clone individually. If the average responses for each J
clone determine a line with no slope in relation to the
abscissa axis, then this reveals that the K spacings are
not different from each other.
These hypotheses for the multivariate profile analysis are
related to three important questions about the data acquired
from Nelder wheel trials:
1. Are the groups (clones) parallel between space points
(different densities)?
2. Are the groups (clones) at equal levels across space
points (different densities)?
3. Do the profiles exhibit flatness across space points
(different densities)?
Based on these null hypotheses, our objectives
were to: i) evaluate eucalypt clones for the relationship
between their tree diameter, height and volume and
their available growing area; ii) analyse autocorrelation
between the observations; and iii) compare the
dendrometric characteristics of diameter, height and
volume between the clones at different densities, using
multivariate profile analysis.
Materials and methods
Study area
The data used come from an experiment with nine eucalypt
clones, distributed in four Nelder wheel plots, located in
the northern state of Tocantins, Brazil (Figure 1). The
experiment was installed in the period from 2010 to 2014 by
the company Eco Brasil Florestas.
The experiment was conducted in a region characterised
by a semi-humid tropical Aw-type climate, according to
Köppen’s classification; high temperatures throughout the
year, with a minimum of 20 °C and a maximum of 32 °C;
a defined rainy season from October to May, and a dry
season from June to September; and annual precipitation of
> 1 800 mm (da Silva Jr 2016).
The predominant vegetation in the region is characteristic
of dense tree savanna, open tree savanna and park
savanna. It presents open xeromorphic vegetation,
dominated and bounded by a herbaceous layer and
grassy–woody carpet, with a heterogeneous floristic
composition (Projeto RadamBrasil 1981).
Description of the experiment
The spacing between the trees along the radius in the
present design is constant; therefore, use of the geometric
progression of the rays ratio (α) is not required in the
estimators. Consequently the spacing area was calculated
using this value multiplied by the increasing Euclidean
distance between two arcs crossing two rays at the
midpoint between trees along theses rays, whose limit is
the side of the triangle formed by their opening and defined
by the angle (θ).
Due to these characteristics, the Nelder wheel plots
were considered an adaptative structure, because, in the
original systematic design, Nelder (1962) proposed that
the experimental arrangement values for the initial radius r0
(distance from the centre of the wheel to the inner border)
should be defined as the ratio for the progression geometry
of the rays (α), the angle between them (θ), and the area
(Ai) per plant.
Nelder wheels have an angle value between the rays
and the different number of rays (on average 96 rays), and
34 concentric arcs (lines) were established in all wheels
(Figures 2 and 3). Information about each Nelder wheel
plot included age, number of rays, the angle between rays,
central geographic coordinates, and the municipality where
Southern Forests 2022, 84(2): 01–12 3

See 3 years
enlarged
area
5 years
6° S 2 years
BRAZIL

Tocantins TOCANTINS
TOCANTIS
SOUTH 4 years
AMERICA

Nelder plots

10° S

0 80 160 km

48° W 44° W

Figure 1: Location of the Nelder wheel sampling plots used in the experiment in the state of Tocantins, Brazil

the experiment was conducted (Table 1). The individual
area per plant was obtained with the following formula:
 De  i 1 De  i 1 _ 
Ai  Di  .   Di De  i
 2 2  (1)
where: Ai is the area per tree; Di is the distance of the trees
between two rays; and D(e)i is the distance between trees
along the radius (approximately 3 m).
The four Nelder wheel sampling plots have different
characteristics for the number of rays and the value of
the opening angle. To minimise these differences, the
spacing treatments were separated into four classes. The
description by classes is shown in Table 2.
The eucalypt clones have different characteristics and
were organised by letters. Two to four different clones were
planted in each Nelder wheel plot of different ages (Table 3).
Data collection
Data were collected in 2016, when the Nelder wheel plots
were two to five years old. These had, on average, 96 rays
and two to four clones, each distributed approximately
between 24 to 40 rays, but the trees were measured only
along 12 rays per clone.
The dendrometric variables measured were: diameter
at breast height (d) at 1.30 m aboveground, and total
height (h). The tree volumes (v) were obtained using the
Schumacher–Hall fitted equation:
ν = 0.000044487 d1.73714816 h1.17608127 (2)
where: v is the total volume per tree (m³); d is the diameter
at breast height (DBH) measured at 1.3 m above ground
(diameter at breast height in cm); and h is the total tree
height (m).
Data analysis
Correlation analysis
The relationship between the variables diameter, height
and volume with the available growing area per tree was
evaluated per clone at different ages, using the correlation
4 Alba da Silva, Netto, Behling, Koehler, Filho and Cerqueira
N between 0.70 and 0.89 considered strong; and from 0.90
0 5 10 20 30 40
trees
Meters
Figure 2: Characteristics of the experimental units of the adapted
Nelder plot sampling circle
Ai
D(e)i
Di
N 0 4 8 16 24 32
Meters
ϴ coincident: there is no effect of the clones.
Figure 3: Characteristics of the experimental units of the adapted
Nelder plot sampling circle. θ = opening angle between the rays;
d(e)i = distance of the trees in the radius (approximately 3 m); di =
distance of the trees between two rays; Ai = area per tree
matrix. The data were subjected to normality evaluation
by the Kolmogorov–Smirnov test at 95% probability. The
variables that did not show normality were transformed
applying the Box–Cox function (1964).
Even using this data transformation technique, the
normality condition was not met; therefore, the Spearman’s
correlation matrix was applied. In addition to this, the
graphical description was used to visualise the behaviour of
the dendrometric variables in the spacing classes.
Siegel (1975) proposed that Spearman correlation
coefficients, which could have positive or negative values,
be classified as follows: between 0 and 0.19 considered a
very weak correlation; between 0.20 and 0.39 considered
weak; between 0.40 and 0.69 considered moderate;
and above considered very strong.
Autocorrelation
The occurrence of autocorrelation and spatial dependence
of the dendrometric variables diameter (d), height (h), and
volume (v) at different ages were verified using graphical
models based on Moran’s statistical index I, using SAS
Studio software with the VARIOGRAM proc function.
The values of this index range from −1 to 1, with positive
ones indicating a positive spatial association between
locations (similarity) and negative ones indicating negative
association (dissimilarity). Values around 0, in envelopes of
complete randomness, indicate random distribution without
spatial association (Moran 1950).
Multivariate profile analysis
Multivariate profile analysis was used to test the difference
between clones in different spacings, once the trees in the
Nelder systematic design might show autocorrelation – a
consequence of non-randomisation and of obtaining data
on an increasing scale of spacing.
The analysis was performed by comparing the clones
in each Nelder wheel plot since the location and age (two,
three, four or five years old) of each one is different. The
response variables were height, diameter and volume.
Systematically, seeking to cover the different spacings,
four samples were selected per clone distributed in each of
the spacing classes. Each sample was composed of three
replicates from the same spacing.
Three hypotheses (H0) of interest are associated and
were tested in the multivariate profile analysis, according to
Rencher (2002):
H0(1) – The average response profiles for different treatments
are parallel: there is no interaction between clones ×
spaces.
H0(2) – The average response profiles for treatments are
H0(3) – The average response profiles for clones are parallel
to the abscissa axis: there is no spacing effect.
Wilk’s lambda test (λ) at 95% probability was used to evaluate
the three hypotheses when analysing three or more clones. This
test is applied by transforming the λ value to a corresponding
F-value and then consulting the F table to evaluate significance.
If the calculated F-value is smaller than the F- value from
the table, the hypotheses H0(1), H0(2) and H0(3) are accepted,
otherwise they are rejected. Whenever tested hypotheses were
rejected, the clones were compared two by two.
When only two clones were present (age two years) they
were evaluated using the Hotelling test (T²) for hypotheses
H0(1) and H0(3) and then comparing the calculated values
with the value in Hotelling’s table (T²). Hypothesis H0(2) was
evaluated using the t-test, as described by Rencher (2002).
SAS Studio software was used with the GLM proc
function. The analysis between two clones was performed
using MATLAB software, in the evaluation version, with
functions programmed by the authors.
Southern Forests 2022, 84(2): 01–12 5

Homogeneity of the variances was verified by the Bartlett
test at 95% probability. Adjustments of the data to the
model’s assumptions were made using log transformation
of the data for the dendrometric variables height, diameter
and volume, when pertinent.
Results
Correlation
Diameter reached the highest correlation values with the
area per tree, ranging from moderate (0.44) to strong (0.88).
Volume was strongly correlated with the available growing
area, whose coefficient values ranged from weak (0.35) to
strong (0.80). Correlation between the variables diameter and
volume with the area per tree was significant for all clones and
showed positive values, indicating a growing trend of these
variables with the increase in the available growing area.
Height showed a very weak to moderate correlation (0.006 to
0.65), with greater correlation at age two years. Clones A, B
and C at four years old, and clone E at three years old resulted
in nonsignificant correlation coefficients.
Autocorrelation
For diameter, height and volume at different ages, the
values of the Moran I index were dispersed in the positive
and negative regions as a function of distance, observed by
Table 1: Information on the Nelder sampling circles of eucalypt
clones used in the present study in the state of Tocantins, Brazil
the trend line, indicating the spatial dependence between
trees, confirming the hypothesis of autocorrelation between
them (Figure 4).
Profile analyses
In general, the eucalypt clones showed an increasing
average of diameter, volume and height with the increase
of the area per tree, with variations in the tree dimensions
in different spacings. Height was the variable that showed
the least variation with the increase in spacing. Table 5
presents the results for the three hypotheses of multivariate
profile analysis for trees of all ages.
Clones H and I: two years old
Clones H and I of Eucalyptus platyphylla, two years old,
showed different characteristics regarding phytosanitary
problems. Tree diameters of the clones at this age did not
show interaction with spacing, and diameter did not differ
significantly among clones. The spacing factor showed a
significant difference, and the highest values were those in
the largest classes, above 11 m² (Figure 5a; Table 5).
All null hypotheses tested for the heights of the clones
were rejected. Therefore, there was interaction between
clones and spacing, and differences between the principal
factors. Clone H showed better results than clone I
(Figure 5b; Table 5). The analysis showed significant
interaction and differences between spacing. The greatest
volumes were obtained at the largest spacing (Figure 5c;
Table 5).

Clones E, G, H and I: three years old

Number Angle Geographic coordinates The results are reported at age three years for clones E and G
Age
of between Municipality of Eucalyptus urograndis, hybrids with different characteristics
(years) S E
rays rays (θ) of productivity and resistance to drought, and for clones H and
2 97 3.71 7°26′29.16″ S 48°05′41.60″ E Araguaína
I of E. platyphylla. All null hypotheses tested for the variables
3 106 3.39 7°04′54.84″ S 48°01′50.79″ E Babaçulândia
diameter and volume were not rejected, meaning that there
4 92 3.91 7°47′40.14″ S 47°30′58.50″ E Barra do Ouro
5 91 3.95 7°06′37.38″ S 48°01′33.65″ E Babaçulândia were no interactions and no differences between the clones
(E, G, H and I) and spacing for plantations three years old
(Figure 6 a,c). At age three years, only the factor ‘clones’
Table 2: Spacing classes defined according to the area per tree, in showed significant differences (Figure 6b).
Nelder’s systematic design experiments in Tocantins, Brazil When comparing the clones two by two, all the differences
were significant, with clone I exhibiting the best results (Table
6; Figure 6b).
Class Description Vital area
I Very dense < 6.0 m²
II Dense 6.0 m² – < 11.0 m² Clones A, B and C: four years old
III Slightly dense 11.0 m² – < 16.0 m² In the Nelder wheel plots at age four years, with clones A, B
IV Not dense ≥ 16.0 m² and C, all E. urograndis hybrids with different characteristics

Table 3: Characteristics of the Eucalyptus spp. clones and information on their inclusion in Nelder wheel plots in Tocantins, Brazil

Clone Characteristics Age (years)

A E. urograndis hybrid
B E. urograndis hybrid
C E. urograndis hybrid
D E. urograndis hybrid
E E. urograndis hybrid
F E. urocam hybrid
G E. urograndis hybrid
H Eucalyptus platyphylla
I Eucalyptus platyphylla
high productivity and medium resistance to drought 4
high productivity, low resistance to drought and phytosanitary problems in low areas 4
high productivity and medium resistance to drought 4
high productivity and low resistance to drought 5
high productivity and medium resistance to drought 3, 5
medium productivity and high resistance to drought 5
high productivity, low resistance to drought and phytosanitary problems in low areas 3
medium productivity, high resistance to drought and phytosanitary problems in low areas 2, 3
medium productivity, high resistance to drought 2, 3
 6 Alba da Silva, Netto, Behling, Koehler, Filho and Cerqueira

2 YEARS
2
1
1
1
MORAN’S I

0
0
0
-1
-1
-1
-2
-2
-2 -1 0 1 2 -3 -2 -1 0 1 2 -2 -1 0 1 2 3
STANDART d STANDART h STANDART v

3 YEARS
1 1
0.5
MORAN’S I

0 0
0.0

-1
-0.5 -1

-2 -1 0 1 -2.0 -1.0 0.0 1.0 -2 -1 0 1 2 3

STANDART d STANDART h STANDART v

4YEARS
1
1 1
MORAN’S I

0
0
0
-1
-1

-2 -1 0 1 -3 -2 -1 0 1 2 -2 -1 0 1
STANDART d STANDART h STANDART v

5 YEARS

1
1
1
MORAN’S 1

0
0
0
-1
-1 -1
-2
-2 -1 0 1 2 -3 -2 -1 0 1 2 -2 -1 0 1 2 3
STANDART d STANDART h STANDART V

Figure 4: Correlograms of diameter, height and volume, at the ages of 2, 3, 4 and 5 years, for eucalypt clones in a systematic Nelder design
located in the state of Tocantins, Brazil
Southern Forests 2022, 84(2): 01–12 7

Table 4: Values of Spearman′s correlation between the while clone C was the best for volume, at spacings of
dendrometric variables height (h), diameter (d), and volume (v), 5–10 m² (Table 7; Figure 7).
with area, for eucalypt clones, at different ages and Nelder circles,
in Tocantins, Brazil
Clones D, E and F: five years old
The results for five-year-old clones used clones D and
Age (years) Clone h (m) d (cm) v (m³) E of E. urograndis hybrids, and E. urocam hybrid F, with
2 H 0.48 * 0.76 * 0.72 * characteristics of medium productivity and high resistance
I 0.65 * 0.82 * 0.80 * to drought. Spacing was the only factor that showed
3 E -0.04 ns 0.53 * 0.41 *
significant differences for the variable diameter. For the
G 0.35 * 0.54 * 0.50 *
variables height and volume the interaction was significant,
H 0.50 * 0.58 * 0.56 *
I 0.42 * 0.78 * 0.72 * while the clones were different for height, and the spacing
4 A 0.006 ns 0.57 * 0.51 * for volume (Figure 8a–c; Table 5).
B 0.12 ns 0.44 * 0.35 * Comparing clones D, E and F for the variable height,
C 0.15 ns 0.47 * 0.39 * only clones E and F were equal, and all other clones were
5 D 0.50 * 0.67 * 0.65 * different (Table 8). Clone D showed the highest height, at
E -0.30 * 0.64 * 0.44 * the spacing 20 m².
F 0.56 * 0.69 * 0.69 * There was an interaction between clones in different
* = significant at 95% probability spacings for the variable height. Clone D showed higher
ns = not significant at 95% of probability values in spacings between 6 m² and 20 m². The three
clones (D, E and F) differed from each other, resulting in
rejection of H0(2) (Table 8). H0(1) was rejected for clones
Table 5: Multivariate analysis for the effect of tree spacing, clone D–E and D–F, indicating the interaction between them in
and interaction (spacing × clone) factors for the dendrometric
different spacings. H0(1) was accepted for clones E and F,
variables height (h), diameter (d), and volume (v), in Nelder wheel
with no interaction between them (Figure 8).
plots, Tocantins, Brazil
The acceptance of H0(2) for the variables diameter and
volume reveals that the clones are similar (i.e. did not differ
Age Tested Calculated statistics Calculated statistically). The clones showed no interaction for the
(years) hypothesis h d v statistics diameter variable in different spacings. Interaction between
2 H0(1) 31.81 * 18.73 ns 35.92 * 22.72 clones and spacings was observed for the variable volume.
H0(2) -6.33 * 1.18 ns -1.78 ns 2.77 H0(3) was rejected, indicating the spacing effect on the
H0(3) 32.34 * 289 * 509 * 22.72 average profile of the diameter and volume variables. Clone
3 H0(1) 0.65 ns 1.25 ns 1.14 ns 1.88 D was the best for the variable volume, at spacing 15–20 m².
H0(2) 4.57 * 0.75 ns 1.13 ns 4.06
H0(3) 0.12 ns 3.44 ns 3.44 ns 3.86
Discussion
4 H0(1) 4.18 * 3.56 * 3.58 * 3.58
H0(2) 6.81 * 5.17 * 6.29 * 5.14
H0(3) 14.45 * 11.35 * 12.7 * 4.75
The highest correlation values were observed for the
5 H0(1) 3.76 * 0.73 ns 8.47 * 3.58
eucalypt clones at the age two years for the three variables,
H0(2) 20.08 * 1.21 ns 1.14 ns 5.14 indicating that at this age the values of diameter, height and
H0(3) 2.52 ns 5.73 * 13.00 * 4.75 volume are more related to spacing than to age. In general,
the clones showed an increasing average behaviour of the
* = significant at 95% probability variables diameter, height and volume with an increase in
ns = not significant at 95% probability the area per tree, with variations in the tree dimensions at
F = tabulated value of the F test at 95% probability different spacings. These results corroborate those found
T² = tabulated value of the Hotelling test at 95% probability by other authors for species of Eucalyptus (Berger et al.
t = tabulated value of the t test at 95% probability 2002; Stape and Binkley 2010; Moulin et al. 2017).
Height was the variable that showed the least variation
with increasing area per tree. This behaviour and the lower
of productivity and resistance to drought, all the null correlation values compared with the variables diameter
hypotheses tested for the variables height, diameter and and volume confirm the density theory of growing trees in
volume were rejected, meaning that there were interactions stands, in which height growth is relatively little affected
and differences between the levels of both principal factors by density (Daniel et al. 1979). The height growth is less
(Figure 7a–c). influenced by spacing and may vary according to the site
When comparing clones A and C, only height differed quality and stand age (Hummel 2000; Kuehne et al. 2013).
between them for the spacing factor. For clones C and B, all In the present study, profile analysis was used to
the comparisons showed differences for all variables, except determine differentiation between eucalypt clones in
for interaction and the principal factor clones for height, to Nelder wheel plots. Thus, the multivariate statistical test of
which the null hypothesis was accepted. For comparisons of profile analysis was effective to overcome the problem of
clones A and B, all the variables differed under all hypotheses. autocorrelation potentially detected in Nelder’s systematic
Clones A, B and C did not differ for the variable diameter design. Effects of clones, density and interaction (clones ×
and presented the highest values at the spacing 20 m². density) on the diameter, volume and height variables were
Clone A presented the highest height, at spacing 5 m², evaluated using profile analysis.
8 Alba da Silva, Netto, Behling, Koehler, Filho and Cerqueira
(a)
13
DIAMETER (cm)
10
7
4 8
5 10 15 20
AREA (m²)
(c) 0.09
0.06
VOLUME (m³)
0.03
5 10 15 20
AREA (m²)
Figure 5: Average profiles for the dendrometric variables diameter, height, and volume in different areas of the eucalypt clones H and I, two years
old, located in the state of Tocantins, Brazil
Autocorrelation between variables in the Nelder wheel
plots is owing to the gradual increase in area per tree,
from the centre to the periphery of the circle. This fact
ensured ordered values of the variables’ growth in the
present study, which were also reinforced by the positive
correlation results of the variables with the area per tree.
Statistically, spatial autocorrelation should be considered
in an evaluation of Nelder wheel trials, because depending
on its distance, measurements on different trees may not
be statistically independent. If we use classical statistical
analyses and the autocorrelation is not considered, it can
lead to erroneous results (Uhl et al. 2015).
According to Fox et al. (2001), a large part of the spatial
dependence observed between trees can be explained by
the development stage of the stand, which depends on
two factors: competition and spatial variation. Young trees
initially suffer less competition and are, therefore, more
influenced by the local effect. As the competition begins,
trees start suffering its effect and the spatial dependence
decreases. After the period of intense competition, natural
mortality provides a reduction in competitive effects
and the attributes of individual trees are once again an
expression of the influences of nearby sites, with spatial
dependence occurring again (Fox et al. 2007). This factor
(b) 17
HEIGHT (m)
14
11
25 5 10 15 20 25
AREA (m²)
H I
25
was observed in this research, with values of Moran’s I
index closer to zero at ages three and four years. At age
five years, Moran’s I index values were higher and similar
to values obtained at age two years.
Corroborating the results of spatial dependence at the
age two years, Teodoro (2016) found spatial dependence
in Nelder wheel plots of Peltophorum dubium (family
Fabaceae) for the variables height, at 6, 13, 25 and 37
months after planting, as well as for diameter, at 13, 25 and
37 months. However, Aquino (2017) did not observe spatial
dependence at age two years in a systematic Nelder’s
design with clones of Eucalyptus urophylla × Eucalyptus
grandis W.Hill ex Maiden. Oda-Souza (2005) found
weak spatial dependence for volume data of Eucalyptus
dunnii Maiden, which would not justify considering it for
conventional statistical analysis.
Since the Nelder wheel plots analysed were for different
ages and with clones of different characteristics, the
analysis was not carried out to compare them, as the
variability in this situation is high. With this experiment
design, multivariate profile analysis provided results of
the variables within each Nelder wheel plot for differences
between clones, differences between spacing, and
interaction between clones and spacing.
Southern Forests 2022, 84(2): 01–12 9

(a) 16 (b) 22

13 18
DIAMETER (cm)

HEIGHT (m)
10 14

7 10

4 6
5 10 15 20 25 5 10 15 20 25
AREA (m²) AREA (m²)
H E G I

0.16
(c)
VOLUME (m³)

0.12

0.08

0.04

5 10 15 20 25
AREA (m²)

Figure 6: Average profiles for the dendrometric variables diameter, height and volume in the different areas for the eucalypt clones E, G, H and I,
for 3-year-old plantations in Tocantins, Brazil

Table 6: Comparison between three-year-old eucalypt clones for the variable height

Source of Calculated statistics Tabulated

variation H–E H–G H–I E–G E–I G–I statistics
Height at age three years
H0(2) −5.88* 3.85* −7.98* 9.06* −9.96* −24.16* 2.77 (t)
* = significant at 95% probability
ns = not significant at 95% probability
t = tabulated value of the t test at 95% probability

Conclusions

Clonal forests are fundamental in locations with greater
water and nutrient restrictions, as in our study region. Nelder
wheel experiments are attractive because of the potential
to represent a wide range of stand densities and growing
conditions, with little demand for space. Multivariate profile
analysis corroborated the desirability of the experimental
project to forest science, because, as pointed out in the
third hypothesis, it proved to be an appropriate and practical
method to avoid the autocorrelation problem in the Nelder
wheel design.
In our study, these allied methodologies suggest that
eucalypt clones with medium to high drought-resistance
characteristics, planted in areas with 6–16 m² per plant, are
recommended for the region where the study was located.
Acknowledgements — We thank the Coordenação de Aperfeiço-
amento de Pessoal de Nível Superior (CAPES) for granting a
master’s scholarship to the first author.
ORCID iDS
Franciele Alba da Silva — https://orcid.org/0000-0002-2054-2531
Sylvio Péllico Netto — https://orcid.org/0000-0002-3178-1810
Alexandre Behling — https://orcid.org/0000-0002-7032-2721
Henrique Koehler — https://orcid.org/0000-0001-7727-7049
Ataídes Marinheski Filho — https://orcid.org/0000-0001-8237-3430
Cláudio Cerqueira — https://orcid.org/0000-0003-1862-7496
10 Alba da Silva, Netto, Behling, Koehler, Filho and Cerqueira

(a) 16 (b) 22

13 18
DIAMETER (cm)

HEIGHT (m)
10 14

7 10

4 6
5 10 15 20 25 5 10 15 20 25
AREA (m²) AREA (m²)
H E G I

0.16
(c)
VOLUME (m³)

0.12

0.08

0.04

5 10 15 20 25
AREA (m²)

Figure 7: Average profiles for the dendrometric variables diameter, height and volume in different areas of eucalypt clones A, B and C, for
4-year-old plantations in Tocantins, Brazil

Table 7: Evaluations of interactions between four-year-old eucalypt Table 8: Comparison of differences between five-year-old eucalypt
clones for the variables height, diameter, and volume clones for the variable height

Calculated statistics Tabulated Calculated statistics Tabulated

Hypothesis Hypothesis
C–A C–B A–B statistics D–E D–F E–F statistics
Height Height
H0(1) 17.724ns 2.442ns 65.954* 22.720 (T²) H0(1) 46.46* 25.87* 19.85ns 22,720 (T²)
H0(2) −0.5764 ns 2.4507ns 3.066* 2.770 (t) H0(2) 6.86* 17.09* −28.27* 2,77 (t)
H0(3) 8.630* 72.941* 19.782* 22.720 (T²) Diameter
Diameter H0(1) 2.05 ns 16.84 ns 5.05 ns 22,720 (T²)
H0(1) 10ns 173.537* 46.7798* 22.720 (T²) H0(2) 2.12 ns −1.12 ns −0.57 ns 2,77 (t)
H0(2) −2ns 9.7423* 8.390* 2.770 (t) H0(3) 99.13* 81.20* 28.96* 22,720 (T²)
H0(3) 5.679ns 442.145* 64.498* 22.720 (T²) Volume
Volume H0(1) 41.79* 18.32 ns 6.16 ns 22,720 (T²)
H0(1) 8.279ns 132.500* 42.430* 22.720 (T²) H0(2) 0.17 ns 1.04 ns −1.58 ns 2,77 (t)
H0(2) −2.345ns 9.060* 9.390* 2.770 (t) H0(3) 0.00005 ns 85.16* 39.44* 22,720 (T²)
H0(3) 5.278ns 539.820* 37.840* 22.720 (T²) * = significant at 95% probability
* = significant at 95% probability ns = not significant at 95% probability
ns = not significant at 95% probability F = tabulated value of the F test at 95% probability
F = tabulated value of the F test at 95% probability T² = tabulated value of the Hotelling test at 95% probability
T² = tabulated value of the Hotelling test at 95% probability t = tabulated value of the t test at 95% probability
t = tabulated value of the t test at 95% probability
Southern Forests 2022, 84(2): 01–12 11

26
(a) 18 (b)

22
15
DIAMETER (cm)

HEIGHT (m)
12 18

9 14

6 10
5 10 15 20 25 5 10 15 20 25
AREA (m²) AREA (m²)
D E F

0.28
(c)
VOLUME (m³)

0.21

0.14

0.07

5 10 15 20 25
AREA (m²)

Figure 8: Average profiles for the dendrometric variables diameter, height and volume in the different areas for the eucalypt clones D, E and F,
for 5-year-old plantations in Tocantins, Brazil

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