Andini 2022 IOP Conf. Ser. Earth Environ. Sci. 951 012091

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3rd International Conference on Agriculture and Bio-industry (ICAGRI 2021) IOP Publishing
IOP Conf. Series: Earth and Environmental Science 951 (2022) 012091 doi:10.1088/1755-1315/951/1/012091
Morphological variation of Aceh Pinus (Pinus merkusii)
R Andini1*, V Melinda1, E Pardede2, L A Yanti1, Khaing Hmon3, R Moulana1, S

Indrioko4
1
Faculty of Agriculture, Forestry Department, Universitas Syiah Kuala, Jalan Tgk.
Hasan Krueng Kalee 3, Darussalam-Banda Aceh 23111, Indonesia
2
Faculty of Agriculture, Universitas HKBP Nommensen, Medan, North Sumatera
3
Khaing Hmon, Harry Perkins Institute of Medical Research, QEII Medical Centre 6
Verdun Street, Nedlands WA 6009, Australia
4
Faculty of Forestry, Gadjah Mada University, Jalan Agro No. 1, Bulaksumur 55281,
Yogyakarta, Indonesia
*Email: rita.andini@unsyiah.ac.id
Abstract. The genus Pinus is classified as one of the most widely distributed genera of conifer
trees, that are majorly distributed in the Northern and Southern hemispheres. Pinus merkusii is
the only member of the genus Pinus that is adapted in the tropics and grows naturally in south
Equator. Its natural distribution found through-out Southeast Asia, e.g. in Indonesia; particularly
on Sumatran Island. Three native strains are known on Sumatra: Aceh, Kerinci, and Tapanuli;
with the first one is known as the largest population among the three and grows naturally on the
high altitudes. The wood has been utilized in building construction, while the resin collected is
classified as one of the most valuable Non Timber Forest Products (NTFP) worth up to US $ 50
million/ year and applied in many industries. Planting of citron grass (Cymbopogon sp.) has been
’booming’ over the past seven years in the middle regions of Aceh due to its attractive selling
price. As consequences, many Pinus trees have been logged and demanded as fire woods in the
distillation process of citron grass. Thus, the citron grass’ plantation has been directly threatening
the existence of Pinus forests. Therefore, in-situ conservation effort should be started from now
on. Here, we would like to report the morphological variation (N= 15 traits) of natural Pinus
(e.g., plant height, diameter of main stem, number of crowns per branch, etc.). The
morphological data collected were then analysed with the Principal Component Analyses (PCA),
in order to cluster the major agronomical parameters among them. This morphological data
would be complemented and benefitted the efforts in the genetic study of Aceh Pinus.
1. Introduction
The genus Pinus or commonly known as ‘pine’ tree in English includes up to 120 species and is
classified as one of the most widely distributed genera of conifer trees. They are majorly distributed in
the northern- and southern hemispheres. In the northern hemispheres, Pinus sylvestris and Pinus eliotti
are accounted as the major components of grown species. They are majorly distributed throughout the
temperate and sub-alpine regions, where they form dense forests of evergreen trees [1]. Meanwhile, P.
merkusii is the only member of the genus Pinus that is native [2] and already adapted in the tropics and
grows naturally in the south equator. There, they can grow up to 80 meters (m) high and can live up to
1,000 years or more under favourable condition. Its natural distribution is found mainly in Southeast
Content from this work may be used under the terms of the Creative Commons Attribution 3.0 licence. Any further distribution
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Asia e.g. in Cambodia, Laos, Myanmar, the Philippines, Thailand, Vietnam, Indonesia, including on the
Sumatran island [3].
On the Sumatran island, three native strains: the Aceh, Kerinci, and Tapanuli pinus grow natively
(see: Figure 1.a.). The three are commonly known as the Sumatran pine or ‘tusam’ tree in bahasa [2].
The Tapanuli strain was initially found in the area of Sipirok in South Tapanuli in 1841 by a german
botanist [4]. Within the Sumatran pine, the Aceh strain is known to be the largest population among the
three and grows naturally and commonly at high altitudes or more than 1,000 m above sea level (a.s.l.)
(Figure 1.b.). Pinus’ wood is being employed in the construction building and classified in grade III and
IV in regard to its strength and long-lasting parameter, respectively. Meanwhile, the collected gum-resin
is classified as one of the local valuable Non Forest Timber Products (NFTP). This is actually a kind of
disintegrated products mainly developed from cellulose decomposition process and produced within the
plant cells. The resin, locally known as ‘gondorukem’ is highly valued in many industries (e.g. in food
industries as water repellent agents, or painting and varnish industries) and worth up to US $ 50 million/
year [5].
Figure 1. a) the map of Indonesia highlighted the distribution of the natural Pinus’ forests on the
Sumatran island in Aceh, Tapanuli, and Kerinci. They are indicated with the red dots (right: map of
was depicted from Imanuddin [6]); b) indicated Pinus’ natural distribution on Sumatra; b) field
location, from which the samples of Aceh pinus was taken. The area was marked as the red circle (left
side); map was developed by Ali M. Muslih (2021).
From the ecological perspective, the existence of Pinus’ trees has been serving as essential indicators
of (i) forest health; particularly those ones located on high altitudes. They have been planted by the
Dutch colonial in the period of 1931-1945 in terms of forest reforestation and rehabilitation programs;
(ii) direct indicator of the ‘severity’ or harshness of the global warming impact, particularly in the
tropics. An increase of the earth’s mean surface temperature by 3.7 oC to 7.8 oC by 2100, this condition
would threaten directly the existence of Pinus’ forests located on the highlands and they would not grow
well, up there. Such condition would not only change the weather patterns; including lead to a more
often occurrence of extreme weather, but this would also cause some dramatical changes in agricultural,
forestry, e.g. shifting land use practices on the tropics, increasing the forest vulnerability status, and
diminishing large areas of forests. If one of these predicted condition happens, other regions situated at
the lower altitudes would be affected adversely and faced many problems related to their agricultural
practiced activities, noted with severe and more often drought periods, etc. [7].
2
A massive deforestation of Pinus’ trees at the lower elevation (below 1,000 m a.s.l.) has been started
over the past two decades. Such environmental disturbance has been triggered due to the ‘booming’ of
citron grass (Cymbopogon sp.) plantation; especially in the middle regions of Aceh. Pinus’ trees have
been logged and demanded as fire woods in the ‘conventional’ distillation process. Farmers on the
highlands would earn an additional income from IDR 110,000 up to 360,000/ l. Such kind of activity
would increase the farmer’s income (Figure 2. a, b, c) [8]. It is being roughly estimated that up to 1.5
m3 volume of wood is required daily in order to complete the boiling process in a big kettle based on a
rapid assessment conducted in 2018. This could be inferred that the plantation of citron grass has been
threatening in-/directly the existence of Pinus’ forests, thus, such massive planting would indirectly
affect the natural balance, and finally also exacerbate the negative impact of global warming particularly
on the high lands that would be predicted to be occurred by 2100 [9].
Figure 2. Pictures in clock-counter wise: a) the Pinus’ forests at the background are competing
with the citron grass plantation on the high lands in Middle Aceh; b) and c) land shifting of former
Pinus’ forests areas into citron grass or other horticultural plants. Such agricultural activities would
not only deteriorate the forests but also the soil condition at the mountainous slopes (pictures belong
one of the authors).
In many cases, the reproductive system in one plant determines indirectly or directly the way genes
are transmitted between generations, and consequently, the levels of genetic variability or diversity are
closely related to this [10]. Biologically, most of conifers are propagated via outcrossing, which is
indicated with a high outcrossing rate and this kind of reproduction mode would directly lead to higher
genetic diversity. Contrastingly, the mode of reproduction within P. merkusii is mainly developed via
self-crossing (selfing). This would resulted a low genetic diversity value of 21% within P. merkusii [5]
based on Simple Sequence Repeat (SSR) marker or microsatellites. Simple Sequence Repeat (SSR) is a
powerful genetic marker technology, which has been developed rapidly over the last decade and is
majorly applied in many biology studies. SSR is referring short stretches of DNA sequence occurring
as tandem repeats of mono-, di-, tri-, tetra-, penta- and hexanucleotides. They are highly polymorphic,
genetic co-dominance, high reproducibility and these properties have provided a number of advantages
over other molecular markers. SSR marker can be applied in a wide range of applications, ranging from
3
genetic mapping, molecular tagging of genes, analyses of genetic diversity within a particular
population, and Marker Assisted Selection (MAS) [11].
Genetic diversity study is essential in order to inform breeder what kinds of basic material they
currently have, that could be further applied in the selection-, or any crop-improvement program that
would support any activities related to the existence of living things on our planet [12]. In adjacent to
that, the genetic diversity assessment can be conducted either in a simple form, namely based on their
morphological characteristics or based on via molecular marker identification. Nevertheless, relying the
characterization of genetic resources based on morphological characteristics or traits solely, are proven
to have a number of limitations, e.g.: a) low polymorphism, b) low heritability, and c) vulnerability to
environmental influences. Therefore, such kind of result should be further confirmed their DNA
variation [13]. These two kinds of assessments (phenotypic and genotypic levels) complement each
other and are essentials in order to develop any important strategies or measured steps related to any
‘further’ conservation efforts, or their efficient uses; particularly in an in-situ conservation program [14];
in this regard related to the in-situ germplasm conservation of Pinus merkusii, that grow naturally in
Middle Aceh regions (see Figure 1.a). Here, we would like to report the morphological variation of
Pinus’ trees grown in a natural forest; such as plant height, diameter of main stem, number of crown per
branch, etc. grown on high altitude (higher than 1.000 m a.s.l.) and low one. The morphological data
collected here is intended to: (1) serve as basic information regarding the still existing Pinus’ forest at
high altitude before they could have been vanished, one day; (2) to classify the main essential sample
based on ‘particular’ morphological characteristics based on a three dimensional scattered plot and the
Principle Component Analyses (PCA).
2. Materials and methods

The field data collection was conducted from June to July 2021. Prior to field assessment, we tried to
2
collect some references for field collection and the morphological traits of Pinus tree [15]. Materials
applied in this study were: i) plastic ruler; ii) measurement band; iii) ‘Haga’ hypso meter; iv) General
Positioning System (GPS), ‘Etrex’ H GPS Garmin (Production Year 2020), and v) writing utensils.
Pictures were taken with a hand-phone camera: Xiaomi DN POCO M3 (Made in China, 2021).
2.1. Assessment of Pinus merkusii’s morphological characteristics

The morphological characteristics of Pinus’ trees grown in natural forest in ‘Rawe’ at an elevation
above 1,000 m. a.s.l. (Figure 3.a) were studied. We decided to take data from the highlands as Pinus’
trees are still available in those locations, but we presumed that they could have been disappeared in two
or three decades from now on if the pace of deforestation could not be halted. The references regarding
the general guidance [2], [16] and environmental condition for Pinus’ habitat were obtained from [17].
There were in total 15 morphological characteristics, e.g. tree height, stem diameter, number of
branches, etc. of each individual tree located at each sub-plot (A, B, C, and D) (Figure 3. b) were studied
based on this reference as general ‘plant descriptor’ [15]. Furthermore, the way we had measured the
morphological data were explained on Table 1. Then, the collected ‘morphological’ data of Aceh Pinus
were averaged and then further analysed with the Principle Component Analyses (PCA). It is a kind of
multivariate analyses in order to classify, group, and finally select the main essential sample based on
‘particular’ or desired characteristics defined by a breeder. The morphological data collected here, would
complement the genetic data of Pinus based on their DNA variation, that would be reserved for another
separate publication, elsewhere.
2.2. Statistical analysis

First, data compilation via excel was made, then the values were averaged. data collected was first noted
in an excel sheet, then the sheet was applied for further analyses e.g., Principle Component Analyses
(PCA) in order to classify the most important morphological characteristics and to enable us to
distinguish the most essential morphological characteristics performed by Pinus’ trees grown on high
4
altitude. The two- and three dimensional PCA is following the previous explanation reported by Andini
et al. [18]
Table 1. The methodology how the morphological characteristics were measured in this study.
No. Morphological Unit Mean of Measurement
Characteristics
A. Qualitative Traits
1. Leaf colour n.a.* Colour based on reference [15]
2. Leaf form n.a. Form based on reference [15]
3. Crown colour n.a. Colour based on reference [15]
4. Fruit colour n.a. Colour based on reference [15]
B. Quantitative Traits [16]
1. Number of plants unit We counted those plants having height higher than 10 m.
per sub-plot located in a sub-plot.
2. Plant height from m Trees that were higher than 10 m. were measured in their
soil height, with the ‘Haga’ hypso meter, year of production in
1990.
3. Height of the first m The height of the first branch from the soil up to the position
branch from soil of the first branch.
4. Main branch’s cm The branch’s diameter was measured, whose height was
diameter at breast approximately 130 cm from the soil surface or approximately
height the height of a person’s breast
5. Stem diameter of cm The stem diameter of the biggest branch, that can be reached
the biggest branch out by a person.
6. Number of unit Manual counting of the maximum number of branches in
branches per tree each tree available
7. Number of crown unit Manual counting of number of crown available at the biggest
per branch branch on each tree.
8. Leaf length cm
9. Needle (leaf) cm Leaf diameter in the form of a ‘needle’ is measured instead of
diameter a measurement band but with a plastic ruler. First, the needle
was flatted with a marker, then the length measured on a
plastic ruler was noticed as diameter or ‘d’ and then, noticed
on the sheet. The diameter was then produced after the value
from manual observation was transformed from diameter (d)
= 2 * radius
10. No. of available unit Manual counting of available seeds
seeds per crown
11. Crown length cm The length of an already fallen, but still ‘intact’, fresh, and fit
in form of a crown was measured with a plastic ruler.
12. Crown diameter cm The diameter was measured with a measurement band.
13. Crown weight g Weighing the weight of a crown with a kitchen scale
*n.a.= not available
5
Location : Rawe (> 1,000 m a.s.l.)

(a) (b) N
1 *
0m
D 2
1
:1
1
gth
3 2
L en
A
1
C 2
1 3
3 B 2
SP= SubPlot 1
1 D *
2
3 1
2
A
C 3
2 3
1 1
B
3 2
SP2
* 1 2
1 A
D 3
2
3
1
1 12
C B 3
2
3
N total= 36 samples SP3
Figure 3. a) the location of research was in Rawe, which is located above 1,000 m a.s.l. and this was
in accordance with Siregar [4], b) visualization how samples were taken from each subplot (SP). Each
SP was then divided into four quadrant regions: A,B,C, and D. From each quadrant, three individuals
were taken as sample in this study and being noted as 1,2,3. From each circle, there individual trees
were taken as sample (N1= 36 samples).
3. Results and discussion

Indonesia is being classified as one of the top-three of the world’s 17 ‘megadiverse’ countries, but
unfortunately, the forested area has been decreased from 62% to 50% in 1990 and 2010, respectively;
according to UN-FAO. Certainly, there are still remain very few large the so called ‘frontier forest’ in
Indonesia, referring to ecologically intact natural forest on the islands of Borneo, Sumatra, and Sulawesi.
However, it is estimated that those ‘frontier forests’ would be depleted until the end of this century, due
to the ‘modernization’ activities that require (massive) land conversion for agricultural production, e.g.
oil palm, rubber, teak plantation or shifting cultivation practices [19] and in this case is the citron grass
plantation in Aceh Tengah that has been competing with Pinus’ forest over the past 7 to 8 years.
The major finding of this study is that the wide eco-geographic distribution of Pinus merkusii; as the
only one in its genera being distributed in the southern hemisphere is here being confirmed. The number
of pine trees is usually increasing in number; particularly in disturbed areas in the tropics with an annual
temperature between 21 up to 28 oC. Its natural habitat is basically ranging from 400 m. up to 2,000 m.
a.s.l. and it is strongly light-demanding [2]. Pinus’ trees are listed as vulnerable 3, therefore, this study
is intended to record the morphological characteristics of those still remaining Pinus’ forests in Aceh
Tengah and targeted for in-situ conservation of local genetic resources.
3.1. Morphological characteristics

Assessment of morphological characteristics is the simplest way to distinguish the genetic variation
among species and this type of assessment has been conducted in many cultivated plants [20]. The
variation of morphological characteristics of Pinus merkusii grown on the elevation above 1,000 m. a.s.l.
6
is summarized on Figure 5 and Table 1. Here, most of the qualitative traits, e.g. leaf, crown, and fruit
colour observed were according to plant descriptor, namely: dark green, gold brown, and yellowish
white, respectively. The leaf appearance of Aceh Pinus was a slightly ‘crooked’ (Figure 4), its form is
classified as simple, spine-shaped leaves3 or also being termed as ‘bifoliar’. Meanwhile, other sub-
tropical Pinus existing having other types of leaf forms referred as monofoliar, trifoliar, and pentafoliar
[17].
Figure 4. Leaf- and crown-morphology of Pinus Aceh (Pinus merkusii) (author’s personal collection).
There were 17 trees per subplot on average and their height was in the range between 17 m to 20 m
(S.D.  0.83) from the soil surface. Trees are commonly standing tall and have a self-supporting growth
form meaning that they can rely on their own supportive tissues rather than a climbing habit to achieve
vertical growth4. The first branch that can be reached out was approximately 3 m on average from the
soil (S.D.  0.73), with main stem diameter ranging from 80 cm to 88 cm (S.D.  2.01). Furthermore,
the stem diameter of the biggest branch was in the range between 10 cm to 22 cm (S.D.  2.49), while
other source mentioned 2-23 cm. [4]. Number of branches per tree was relatively high, namely ranging
from 75 to 98 branches (S.D.  5.03). Similarly, each tree possesses relatively high number of crown or
also referred as ‘cone’ sometimes, in which 43 up to 96 crowns were counted (S.D.  13.54) (Figure 5).
Crown is generally defined as seed-bearing organ on many gymnosperm plants. A type of fruit,
usually, woody, ovoid to globular, including scales, bracts, or bracteoles arranged around a central axis,
especially found in confiners and cycads3. Moreover, each crown has 56 seeds on average (S.D.  3.09),
although up to 65 seeds can be borne in one crown. Such high seeds’ bearing capacity might allow many
conifers’ plant to conduct outcrossing and at the end to broaden the genetic variation pattern [21].
Although, self-pollination in some cases would be also possible, such as by the case of P. merkusii
grown in natural forests of Thailand [3]. Crown length was in the range between 8 cm to 9 cm (S.D. 
0.49), and a diameter between 4 cm to 5 cm (S.D.  0.27) (Figure 5).
Our current findings in terms of morphological characteristics of P. merkusii are in accordance with
an earlier report published by Siregar [4]. The height of trees was in the range between 20 m. up to 40
m., bifoliar leaf type, leaf appearance resembles to that of needles with scaly on the surface. Moreover,
male and female flowers existing in one plant. Male flowers are approximately 2 cm. long located at the
entry point of young shoot. Meanwhile, the female ones are clustered in a small number on the tip of
young shoot, found in a slightly crooked cylindrical form. The length of crown is in line with our current
findings, namely in the range between 7 cm. to 10 cm. and the crown usually bears high number of seeds
and weights approximately 25 g. each. Length of seed was varied from 6-7 mm. The colour of the main
stem’s bark is generally dark brown and up to 100 cm. of stem diameter has been recorded. Meanwhile,
the diameter of side branches’ is 20% smaller or between 2 to 23 cm. on average (Figure 5). These
values measured from the natural populations grown in south Tapanuli are also in line with our current
results, here. Similar report highlighted that the main stem of Pinus’ strain grown in Aceh is less straight
7
compared to the other strains (Tapanuli and Kerinci) and this stem straightness could be also applied as
one of major breeding targets other than high resin yield. In this study, resin amount of 80 -166 g per
tree per three days was noted and this value is also in accordance with the previous result published by
Susilowati and Rachmat [22].
B.1. B.2. B.3.
15 16 17 18 19 20
No.of plants h. > 10 m. per SubPlot (unit) Plant height from soil (m) Height of the first branch from soil (cm)
B.4. B.5. B.6.
Main brach’s diameter at breast height (cm) Stem diameter of the biggest branch (cm) Number of branches per tree (unit)
B.7. B.8. B.10.
No. of crown per branch (unit) Leaf length (cm) No. of available seeds per crown (unit)
B.11. B.12. B.13.
Crown length (cm) Crown diameter (cm) Crown weight (g)
Figure 5. Histogram distribution of quantitative morphological characteristics of P. merkusii grown on

the natural habitat in Aceh (developed by JMP version 16.0, SAS Inc., year 2021, university license);
explanation of B.1., B.2., B.3., etc., see: Table 1.
3.2. Principle Component Analyses (PCA)

Based on a three-dimensional scattered plot, thus, the total number of samples is generally clustered
into three categories (Figure 6). The first cluster was grouping those accessions (accessions no. 13, 14,
15, …., 24) having a strong tendency to be more superior in : crown characteristics, higher seed numbers
per crown, and number of plants higher than 10 m. per plot (indicated in red colour). They were found
majorly on subplot 2. Meanwhile, the subplot 1 consisting of accessions’ no. 1, 2, 3, …, 12 were more
superior in terms of plant height, branch-, and the crown length. Those accessions located on subplot 1
might have received more sun-light intensity; compared to the other locations or subplots as higher sun-
light intensity might promote better plant growth and elongation 4. The last cluster was indicated with
green colour and consisted of accessions belong to subplot 3, namely: accessions’ no. 25, 26, 27, …, 36.
8
Figure 6. A three-dimensional scattered plot of the morphological characteristics of Aceh Pinus based
on quantitative traits (developed by JMP version 16.0, SAS Inc., edition year 2021).
Figure 7. Principle Component Analyses (PCA) of Aceh Pinus based on correlation, which were
comprised from 15 morphological characteristics; mostly from the quantitative ones (developed by
JMP version 16.0, SAS Inc., edition year 2021).
Principle Component Analyses (PCA) is majorly a data reduction technique in order to simplify
many parameters as there are in total 15 morphological characteristics being analysed. Here, we
confirmed that PCA shown in Figure 7 was proven to be effective to differentiate the three cluster
similarly to the previous figure (Figure 6). PC 1 explains 19.1 % of existing variations, those are linked
with plant height, branch height, and the crown length. In adjacent to that, PC 2 explains only 16.1 of
variations that are closely related with the major characteristics of crown e.g., number of crowns per
branch, crown diameter, crown and leaf length. Here, we can imply that those are closely related with
9
reproductive ones. Furthermore, we can also conclude that almost all morphological characteristics are
the same important as this is indicated with relatively low number of percentages of each PC axis (Figure
7). Breeder should select accession no. 25 for individual attributed with many existing crowns per
branch, no. 6 for plant height, and no. 19 for individual with an extraordinary crown diameter.
4. Conclusions
The morphological characteristics of Aceh Pinus (Pinus merkusii) strain were studied, here. There were
in total 17 traits, in which four were representing the qualitative ones, e.g. leaf color, leaf form, crown
color, and fruit color. Plant height was varied between 17-20 m. from the soil surface, stem diameter
was 80 – 88 cm., stem diameter of the biggest branch was 10 – 22 cm. Number of branches per tree were
75 – 98 branches, in which 43 to 96 crowns were available per branch. Each crown possesses 56 or
sometimes up to 65 seeds that allow pine tree to reproduce the whole year. Principle Component
Analyses (PCA) and three-dimensional scatter plot enabled us to distinguish each accession based on
three major groups or clusters. Breeder should select accession no. 25 for individual attributed with
many existing crowns per branch, no. 6 for plant height, and no. 19 for individual with an extraordinary
crown diameter.
Acknowledgements
The research was financed by LP2M Universitas Syiah Kuala (USK), Ministry of Education and Culture,
based on the contract document for ‘Lektor’ scheme, No. 172/UN11/SPK/PNBP/2021 dated on 22 nd
February 2021. Author would like to express our genuine gratitude to Mr. Ali M. Muslih from the
Forestry Department for providing us a map of the field data collection. Last but not least, we would
like to express our sincere appreciation to those who assisted us during the field work, and especially to
Dr. Joko R. Withono, and Dr. Izu Andry (Kebun Raya Bogor), and Dr. Revis Asra from Universitas
Andalas, Sumatra Barat for the later analyses to complement the genetic diversity study.
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11

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This content was downloaded from IP address 103.94.190.23 on 19/04/2024 at 09:21

Morphological variation of Aceh Pinus (Pinus merkusii)

R Andini1*, V Melinda1, E Pardede2, L A Yanti1, Khaing Hmon3, R Moulana1, S

2. Materials and methods

2.1. Assessment of Pinus merkusii’s morphological characteristics

2.2. Statistical analysis

Location : Rawe (> 1,000 m a.s.l.)

N total= 36 samples SP3

3. Results and discussion

3.1. Morphological characteristics

B.1. B.2. B.3.

B.4. B.5. B.6.

B.7. B.8. B.10.

B.11. B.12. B.13.

Crown length (cm) Crown diameter (cm) Crown weight (g)

Figure 5. Histogram distribution of quantitative morphological characteristics of P. merkusii grown on

3.2. Principle Component Analyses (PCA)

Antirrhinum Ann. Bot. 98 1053–60

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