Nikolić et al., 2019, Biologica Nyssana 10(1)


10 (1) September 2019: 9-16
DOI: 10.5281/zenodo.3463982

Morphometric analysis of 
vegetative and reproductive 
organs of the Fragaria species

Original Article

Danijela Nikolić
Department of Biology and Ecology, Faculty of 
Sciences and Mathematics, University of Niš, Serbia
danid@pmf.ni.ac.rs (corresponding author)

Milena Veličković
Department of Biology and Ecology, Faculty of 
Sciences and Mathematics, University of Niš, Serbia
milenavelickovic9890@gmail.com

Irena Raca
Department of Biology and Ecology, Faculty of 
Sciences and Mathematics, University of Niš, Serbia
racairena@gmail.com

Dragana Jenačković Gocić
Department of Biology and Ecology, Faculty of 
Sciences and Mathematics, University of Niš, Serbia
draganaj@pmf.ni.ac.rs

Marina Jušković
Department of Biology and Ecology, Faculty of 
Sciences and Mathematics, University of Niš, Serbia
marinaju@pmf.ni.ac.rs

Vladimir Ranđelović
Department of Biology and Ecology, Faculty of 
Sciences and Mathematics, University of Niš, Serbia
vladar@pmf.ni.ac.rs

Received: July 29, 2019
Revised: September 9, 2019
Accepted: September10, 2019

Abstract: 
This paper presents results of a study on morphological variability of Fra-
garia vesca, F. moschata and F. viridis populations collected in their natural 
habitats in Serbia, Macedonia and Montenegro. The goals of the study were 
to determine the degree of morphological variability as well as the morpho-
logical characters with the most significant impact on differentiation of these 
species. The morphometric study was carried out on 7 populations (142 indi-
viduals x 19 morphological characters). Analyses included standard univari-
ate (Descriptive, ANOVA) and multivariate (Principal Component Analysis, 
Canonical Discriminant Analysis, UPGMA classification) analyses. The ob-
tained results indicate moderate to high variability in most of the analysed 
characters. Fragaria moschata has shown the highest level of variability 
for almost all characters in comparison to F. vesca and F. viridis. The most 
significant differentiation in Fragaria species was present in following char-
acters: Leaf-L, Lam-cent-L, Lam-cent-W, Lam-lat-L, Ped-L and Sep-int-L.
Key words: 
Fragaria, morphological characters, variability pattern

Apstract: 
Morfometrijska analiza vegetativnih i reproduktivnih organa vrsta roda 
Fragaria
U ovoj studiji istraživana je morfološka varijabilnost populacija vrsta Fra-
garia vesca, F. moschata i F. viridis sakupljenih na njihovim prirodnim 
staništima u Srbiji, Makedoniji i Crnoj Gori. Ciljevi ovog rada su bili da se 
ustanovi nivo morfološke varijabilnosti kao i da se utvrde morfološki ka-
rakteri koji najviše doprinose u diferencijaciji ovih vrsta. Morfometrijska 
studija je sprovedena na 7 populacija (142 individua x 19 morofoških ka-
raktera). Urađene su standardne statističke metode univarijantne (Deskripi-
tivna, ANOVA) i multivarijantne (Analiza glavnih komponenti (PCA), 
Kanonijska diskriminantna analiza (CDA), UPGMA klasifikacija) anal-
ize. Dobijeni rezultati ukazuju na umerenu do visoku varijabilnost većine 
analiziranih karaktera. Fragaria moschata je pokazala najveći stepen vari-
jabilnosti za skoro sve karaktere u poređenju sa F. vesca i F. viridis. Kara-
kteri koji su najviše doprineli u diferencijaciji istraživanih Fragaria vrsta 
su: Leaf-L, Lam-cent-L, Lam-cent-W, Lam-lat-L, Ped-L and Sep-int-L.
Ključne reči: 
Fragaria, morfološki karakteri, varijabilnost

Introduction

Genus Fragaria L. includes 24 species (DiMeglio et 
al., 2014) distributed in Eurasia, North America and 
South America. Flora Europea (Tutin et al., 1968) 
lists 5 species: Fragaria vesca L., F. moschata We-
ston, F. viridis Weston, F. virginiana Mill. and F. Í 
ananassa (Weston) Rozier, while Euro+Med Plant-
base also includes F. chiloensis (L.) Weston.

In Serbia, this genus is represented with three wild 

species: F. vesca, F. moschata and F. viridis (Gajić, 
1972). These species are also present in Macedonia 
and Montenegro (Micevski, 1998; Euro+Med Plant-
base). Fragaria vesca has the Eurasian origin and 
the most extensive native range among all Fragaria 
species, as it is distributed from the west of the Urals 
throughout Northern Europe and across the North 
American continent (Hummer et al., 2011). In Serbia 
it has a wide distribution (Gajić, 1972) and may be 
found in meadows and forests in following vegeta-

© 2019 Nikolić et al. This is an open-access article distributed under the terms of the Creative Commons 
Attribution License, which permits unrestricted use, distribution, and build upon your work non-commercially 
under the same license as the original. 9



tion associations: Quercetum frainetto-cerris, Quer-
ceto-Carpinetum, Quercetum montanum, Fagetum 
montanum.

Fragaria moschata is distributed in Central and 
Southern Europe and the central part of Russia and 
Ukraine. In Serbia this species grows in forests 
(Querceto-Fraxinetum, Quercetum farnetto-cerris 
and Querceto-Carpinetum), at forest edges, burned 
areas, open grassy hillsides and meadows, usually in 
shadowy places and tall grass, preferring saturated 
fertile soils.

Fragaria viridis is distributed almost throughout 
Europe, as well as in Siberia and at the Canary Is-
lands. In Serbia it is recorded at meadows (Galieto-
Festucetum vallesiacae, Chrysopogonetum pannon-
icum), forests and burned areas (Gajić, 1972).

Morphological recognition among species is 
complicated within the genus Fragaria, due to high 
levels of morphological variations and similarities 
(Nosrati et al., 2015). 

Diploids, tetraploids, hexaploids, octaploids and 
decaploids have been reported, as well as several 
intermediate ploidy levels of interspecific hybrids 
of Fragaria species (Nosrati et al., 2015). Fragar-
ia vesca and F. viridis are diploids, while F. mos-
chata is the only hexaploid species (DiMeglio et al., 
2014). The best-studied form is F. vesca, included in 
many studies dealing with morphological variability 
caused by ecological conditions (Hancock & Bring-
hurst, 1978; Jenson & Hancock, 1981; Huseinović 
& Osmanović, 2010; Malinikova et al., 2013; 
Huseinović et al., 2014) or by genetic diversity 

Fig 1. Some morphological characters of the vegetative and reproductive organs of the analysed species: A-Fragaria 
moschata, B-F. vesca, C-F. viridis (1-Leaf-L, 2-Peti-L, 3-Lam-cent-L, 4-Lam-cent-W, 5-Lam-lat-L, 6-Lam-lat-W, 7-Lob-
lam-cent-N, 8-Scap-L, 9-Ped-L, 10-Sep-ext-L, 11-Sep-ext-W, 12-Sep-int-L, 13-Sep-int-W, 14-Pet-L, 15-Pet-W).

(Labokas & Bagdonaitë, 2005; Zhang et al., 2014). 
The quantification of variation at morphological and 
molecular levels within and between populations 
of F. vesca and the American octoploid species F. 
chiloensis and F. virginiana have been described by 
Harisson et al. (1997, 2000). Phenotypic variation 
within the diploid section of genus Fragaria was 
also a subject of several studies (Sargent et al., 2004; 
Labokas & Bagdonaitë, 2005). However, no quanti-
tative studies of morphological variability have been 
carried out to measure the phenotypic variation be-
tween diploid and hexaploid Fragaria species. 

This study aimed to research the morphological 
variability of vegetative and reproductive organs of 
two diploid Fragaria species (F. vesca and F. viridis) 
and a hexaploid species F. moschata and to deter-
mine the morphological characters with the most 
significant impact on differentiation between these 
species.

Materials and methods
Sample collections

The seven populations of Fragaria species (F. vesca, 
F. viridis and F. moschata) were collected for mor-
phological analysis. The samples were taken from 
Serbia, Montenegro and Macedonia (Tab. 1) dur-
ing two vegetation seasons (2017 and 2018). The 
voucher specimens were deposited in the Herbarium 
Moesiacum Niš (HMN) at Department of Biology 
and Ecology, University of Niš. Identification of col-
lected material was made by using literature sources 

10

BIOLOGICA NYSSANA ● 10 (1) September 2019: 9-16 Nikolić et al. ● Morphometric analysis of vegetative and reproductive 
organs of the Fragaria species



(Tutin et al., 1968; Gajić, 1972; Markova, 1973).
Morphometric analyses
The 7 morphological characters of vegetative or-
gans and 12 morphological characters of reproduc-
tive organs were selected for analyses. Vegetative 
morphological characters included: length of leaf 
(Leaf-L ), length of petioles (Peti-L), length of cen-
tral lamina (Lam-cent-L), width of central lamina 
(Lam-cent-W), number of lobes on central lamina 
(Lob-lam-cent-N), length of lateral lamina (Lam-
lat-L) and width of lateral lamina (Lam-lat-W). The 
reproductive morphological characters included: 
length of scape (Scap-L), length of peduncles (Ped-
L), length of external sepal (Sep-ext-L), width of 
external sepal (Sep-ext-W), number of lobes on 
external sepal (Lob-sep-ext-N), number of external 
sepals with lobes (Sep-ext-Lob-N), length of inter-
nal sepal (Sep-int-L), width of internal sepal (Sep-
int-W), length of petal (Pet-L), width of petal (Pet-
W), number of flowers (Flow-N), number of flower 
branches (Flow-branch-N) (Fig. 1).

The plant material was first digitised by the cam-
era (Sony 20 megapixels) and then measured using 
ImageJ software.
Statistical analysis
Descriptive statistics (mean, max, min, standard de-
viation and coefficient of variation) were computed 
to evaluate the range of variation for morphological 
traits.

The coefficient of variability was used to describe 

BIOLOGICA NYSSANA ● 10 (1) September 2019: 9-16 Nikolić et al. ● Morphometric analysis of vegetative and reproductive 
organs of the Fragaria species

11

the morphological variability of characters. Moder-
ately variable characters are those with CV=20-40%, 
highly morphologically variable had CV higher than 
40%, while stable characters are those with a coef-
ficient of variation below 10%.

Analysis of variance (ANOVA) was performed 
to identify levels of significant variation for each 
character. Multivariate analyses (Principal compo-
nent analysis (PCA), Canonical discriminant analy-
sis (CDA) and Cluster analyses (UPGMA) based on 
Mahalanobis distances) were performed to examine 
relationships between the studied species and to 
find the best characters that facilitate discrimina-
tion between the species. Statistical analyses were 
performed using the package Statistika 8.0 (Statsoft 
2007). 

Results and discussion
Morphological variability of vegetative and 
reproductive characters - descriptive analysis

The results of the descriptive statistics have shown 
significant variability for almost all morphological 
characters (Tab. 2) 

Generally, characters of vegetative and reproduc-
tive organs of Fragaria species have shown moder-
ate to high degrees of variability. Following char-
acters may be considered most variable within F. 
moschata populations, characterised by large-scale 
deviation of maximum and minimum values from 
the mean values, and with highest standard devia-
tion values: Ped-L (23.40 ± 9.01), Leaf-L (22.55 ± 

No. Taxon Locality Number of 
individuals

Voucher 
number 
(HMN)

Legators

1. Fragaria viridis Stara planina 
Mt. (Serbia)

20 13697 Veličković, M.
Nikolić, D.

2. Fragaria moschata Rtanj Mt. 
(Serbia)

20 13703 Veličković, M.
Nikolić, D.

4. Fragaria moschata Vlasina (Serbia) 20 13701 Veličković, M.
Nikolić, D.

3. Fragaria vesca Stol Mt. (Serbia) 20 13707 Veličković, M.
Nikolić, D.

5. Fragaria vesca Kraljevica 
(Serbia)

12 13710 Veličković, M. 
Nikolić, D.

6. Fragaria vesca Jablanica
(Macedonia)

20 13958 Raca I.
Ranđelović, V.

7. Fragaria vesca Prokletije
(Montenegro)

30 13959 Raca, I.,
Ljubisavljević, I. 
Jenačković, D.
Ranđelović, V.

Table 1. The list of analysed populations of the Fragaria species



12

BIOLOGICA NYSSANA ● 10 (1) September 2019: 9-16 Nikolić et al. ● Morphometric analysis of vegetative and reproductive 
organs of the Fragaria species

 
 F

ra
ga

ria
 v

es
ca

 
Fr

ag
ar

ia
 m

os
ch

at
a 

Fr
ag

ar
ia

 v
iri

di
s 

C
ha

ra
ct

er
N

M
ea

n
M

in
.

M
ax

.
St

d.
D

ev
.

C
V 

%
 N

M
ea

n
M

in
.

M
ax

.
St

d.
D

ev
.

C
V 

%
 N

M
ea

n
M

in
.

M
ax

.
St

d.
D

ev
.

C
V 

%

Le
af

-L
 (c

m
)

82
12

.0
2

4.
83

22
.6

9
3.

91
32

.5
3

35
22

.5
5

8.
67

39
.9

6
7.

06
31

.2
9

20
11

.5
0

6.
09

16
.7

3
2.

54
22

.0
4

Pe
ti-

L 
(c

m
)

82
8.

39
2.

60
16

.7
7

3.
22

38
.4

2
35

16
.4

9
5.

40
28

.8
3

5.
72

34
.7

0
20

8.
29

3.
06

12
.7

1
2.

24
27

.0
1

La
m

-c
en

t-L
 (c

m
)

82
3.

34
1.

99
5.

06
0.

76
22

.6
7

35
5.

32
2.

68
10

.4
9

1.
62

30
.5

5
20

2.
90

2.
02

4.
02

0.
66

22
.9

0

La
m

-c
en

t-W
 (c

m
)

82
2.

47
1.

31
3.

81
0.

53
21

.4
3

35
3.

44
1.

74
6.

52
0.

93
26

.9
4

20
2.

06
1.

19
2.

86
0.

47
22

.9
2

Lo
b-

 la
m

-c
en

t-N
82

7.
44

5.
00

11
.0

0
1.

32
17

.6
9

35
7.

86
6.

00
10

.0
0

1.
06

13
.5

1
20

7.
60

6.
00

10
.0

0
1.

19
15

.6
3

La
m

-la
t-L

 (c
m

)
82

2.
74

1.
37

4.
91

0.
86

31
.5

5
35

4.
48

2.
42

8.
00

1.
16

25
.8

5
20

2.
74

1.
81

3.
76

0.
54

19
.7

9

La
m

-la
t-W

 (c
m

)
82

2.
19

1.
06

3.
90

0.
63

30
.5

7
35

3.
03

1.
58

5.
06

0.
81

26
.7

6
20

1.
92

1.
37

2.
70

0.
40

20
.6

1

Sc
ap

-L
 (c

m
)

81
2.

34
0.

70
10

.1
1

1.
70

72
.7

6
35

4.
25

1.
31

8.
60

2.
09

49
.2

2
20

3.
00

1.
35

6.
00

1.
13

37
.5

9

Pe
d-

L 
(c

m
)

81
11

.9
7

4.
06

20
.5

5
3.

72
31

.1
0

35
23

.4
0

9.
16

41
.0

6
9.

01
38

.5
2

20
9.

22
6.

16
13

.2
1

1.
79

19
.4

5

Se
p-

ex
t- 

L 
(c

m
)

82
0.

40
0.

23
0.

80
0.

10
24

.9
5

34
0.

49
0.

25
0.

91
0.

15
31

.3
7

20
0.

28
0.

21
0.

42
0.

06
21

.5
4

Se
p-

ex
t-W

 (c
m

)
82

0.
13

0.
06

0.
23

0.
04

29
.4

5
34

0.
11

0.
04

0.
19

0.
04

39
.6

7
20

0.
08

0.
06

0.
13

0.
02

21
.1

4

Lo
b-

se
p-

ex
t-N

73
2.

00
2.

00
2.

00
0.

00
0.

00
22

2.
00

2.
00

2.
00

0.
00

0.
00

20
2.

00
2.

00
2.

00
0.

00
0.

00

Se
p-

ex
t-l

ob
-N

43
1.

37
1.

00
3.

00
0.

62
45

.0
5

22
1.

45
1.

00
5.

00
0.

91
62

.6
8

20
1.

55
1.

00
2.

00
0.

51
32

.9
3

Se
p-

in
t-L

 (c
m

)
82

0.
42

0.
26

0.
71

0.
08

20
.3

5
35

0.
59

0.
26

0.
97

0.
17

27
.9

6
20

0.
33

0.
26

0.
45

0.
06

18
.7

4

Se
p-

in
t-W

 (c
m

)
82

0.
19

0.
09

0.
40

0.
05

24
.9

9
35

0.
19

0.
11

0.
38

0.
05

26
.6

6
20

0.
14

0.
10

0.
21

0.
03

22
.8

6

Pe
t-L

 (c
m

)
71

0.
49

0.
28

0.
88

0.
14

28
.8

8
34

0.
68

0.
29

0.
94

0.
17

25
.1

0
20

0.
38

0.
25

0.
66

0.
12

32
.0

0

Pe
t-W

 (c
m

)
71

0.
48

0.
21

0.
90

0.
16

32
.8

2
34

0.
58

0.
24

1.
00

0.
17

29
.9

8
20

0.
32

0.
18

0.
50

0.
08

25
.7

6

Fl
ow

-N
82

3.
15

1.
00

6.
00

1.
33

42
.1

2
35

4.
09

1.
00

8.
00

1.
67

40
.8

6
20

2.
50

1.
00

5.
00

1.
05

42
.0

5

Fl
ow

-b
ra

nc
h-

N
82

1.
40

1.
00

4.
00

0.
66

47
.3

5
35

1.
00

1.
00

1.
00

0.
00

0.
00

20
1.

50
1.

00
3.

00
0.

61
40

.4
7

Ta
bl

e 
2.

 R
es

ul
ts

 o
f d

es
cr

ip
tiv

e 
st

at
is

tic
s 

fo
r t

he
 m

or
ph

ol
og

ic
al

 c
ha

ra
ct

er
s 

of
 th

e 
Fr

ag
ar

ia
 s

pe
ci

es
.



13

7.06), Peti-L (16.49 ± 5.72). The number of lobes 
on central lamina (Lob-lam-cent-N) ranged from 6 
to 10 lobes. The least variable character of the veg-
etative region within F. moschata populations was 
Lam-lat-W (3.03 ± 0.81). Of characters of the flow-
ering parts, characters Sep-ext-W (0.11 ± 0.04) and 
Sep-int-W (0.19 ± 0.05) were distinguished as the 
least variable. The number of lobes on external sepal 
(Lob-sep-ext-N) was 2, while number of external se-
pals with lobes (Sep-ext-Lob-N) ranged from 1 to 5. 
The number of flowers (Flow-N) was represented by 
a range from 1 to 8, while number of flower branch-
es (Flow-branch-N) was 1. 

Slightly smaller variability of characters was re-
corded within F. vesca populations in comparison to 
F. moschata populations. The most variable charac-
ters of the vegetative region were Leaf-L (12.02 ± 
3.91) and Peti-L (2.60 ± 16.77). The number of lobes 
on the central lamina (Lob-lam-cent-N) ranged from 
5 to 11. The least variable characters of the vegeta-
tive region in F. vesca populations were Lam-cent-W 
(2.47 ± 0.53) and Lam-lat-W (2.19 ±0.63). Among 
the characters of the flowering parts, characters Sep-
ext-W (0.13 ± 0.04), Sep-int-W (0.19 ± 0.05) and 

Sep-int- L (0.42 ± 0.08) were distinguished as the 
least variable.

Lobes on external sepals (Lob-sep-ext-N) was 
2, while external sepals with lobes (Sep-ext-Lob-N) 
ranged from 1 to 3. Flowers (Flow-N) was from 1 
to 6, while flower branches (Flow-branch-N) ranged 
from 1 to 4. 

The morphological characters of vegetative and 
reproductive organs of F. viridis individuals have 
shown the least variability when compared with F. 
moschata and F. vesca populations. The most vari-
able characters of the vegetative region were Leaf-L 
(11.5 ± 2.54) and Peti-L (8.29 ± 2.24). The number 
of lobes on central lamina (Lob-lam-cent-N) ranged 
from 6 to 10.

The least variable characters of the vegetative re-
gion within F. viridis populations were Lam-lat-W 
(1.92 ±0.40) and Lam-cent-W (2.06 ± 0.47). Within 
the characters of the flowering parts, characters such 
as Sep-ext-W (0.08 ± 0.02) and Sep-int-W (0.14 ± 
0.03) were determined to be the least variable.

Lobes on external sepals (Lob-sep-ext-N) was 
2, while external sepals with lobes (Sep-ext-Lob-N) 
ranged from 1 to 2. Flowers (Flow-N) was repre-

BIOLOGICA NYSSANA ● 10 (1) September 2019: 9-16 Nikolić et al. ● Morphometric analysis of vegetative and reproductive 
organs of the Fragaria species

Acronyms F p PCA 1 PCA 2 PCA 3
Leaf-L 64.718 0.000 -0.932 -0.160 -0.080
Peti-L 56.213 0.000 -0.898 -0.129 -0.104
Lam-cent-L 52.904 0.000 -0.919 -0.173 0.066
Lam-cent-W 37.295 0.000 -0.862 -0.150 0.176
Lam-lat-L 47.633 0.000 -0.907 -0.257 -0.116
Lam-lat-W 31.539 0.000 -0.873 -0.230 -0.012
Lam-cent- lob-N 1.407 0.249 -0.398 -0.092 0.225
Scap-L 65.050 0.000 -0.898 -0.131 -0.028
Sep-Int-L 45.775 0.000 -0.672 0.477 -0.137
Sep-Int-W 8.180 0.000 -0.194 0.644 0.334
Pet-L 29.814 0.000 -0.616 0.450 -0.226
Pet-W 17.237 0.000 -0.499 0.605 -0.072
Sep-Ext-L 21.148 0.000 -0.578 0.552 -0.147
Sep-Ext-W 14.970 0.000 -0.079 0.701 0.378
Lob- Sep-Ext-N 11.130 0.000 0.398 0.413 -0.520
Flow-branch-N 7.503 0.001 0.407 0.050 0.328
Flow-N 9.115 0.000 -0.427 0.246 0.488
Ped-L 15.185 0.000 -0.573 -0.202 -0.301
Sep-Ext-Lob-N 3.837 0.025 0.234 0.263 -0.661

Table 3. Results of ANOVA and PCA analysis for quantitative characters of individuals of the 
Fragaria species.

(F- Fisher’s coefficient, p- the level of significance (p<0.05); PCA axes- Component 
with loadings >0.700 is highlighted in bold)



sented by a range from 1 to 5, while flower branches 
(Flow-branch-N) ranged from 1 to 3. 

According to values of the coefficient of varia-
tion, the most variable character (CV>70%) within 
the F. vesca populations was Scap-L. High level 
of variability was shown by characters Sep-ext-
lob-N, Flow-N and Flow-branch-N. One character 
had no detected variability: Lob-sep-ext-N. All the 
other characters have shown moderate variability 
(20%<CV>40%). Morphological characters with 
high variability within F. moschata populations 
were: Sep-ext-lob-N, Scap-L, and Flow-N. All the 
other characters have shown moderate variability, 
except character Lob-lam-cent-N with low variabil-
ity and Lob-sep-ext-N and Flow-branch-N, which 
did not show any variability (Tab. 2). The characters 

with low variability (CV<20%) within population 
of F. viridis were Lob-lam-cent-N, Lam-lat-L, Ped-
L and Sep-int-L. Only Flow-N and Flow-branch-N 
have shown high variability (CV>40%), while all 
other investigated characters have shown a moder-
ate degree of variability (Tab. 2).
Analysis of variance (ANOVA)
The ANOVA test has shown that all characters, ex-
cept Lam-cent-lob-N, were statistically significant 
(p< 0.005) (Tab. 3). According to Fisher’s coef-
ficient (F), almost all vegetative characters may be 
treated as the most significant (F>30). Among the 

characters of flowering region, Scap-L (F = 65.05) 
and Sep-Int-L (F = 45.78) were the most significant. 
This set of characters with the highest F values is 
the most suitable for describing morphological dif-
ferentiation between Fragaria species, based on the 
results of this analysis.

Morphological differentiation of analysed 
Fragaria species 

The first three principal components accounted for 
57.44% of the total observed variance (Tab. 2). The 
characters which contributed the most to variabil-
ity along the first axis (36.19%) are the following: 
Leaf-L, Lam-cent-L, Lam-cent-W, Lam-lat-L, Ped-
L and Sep-int-L. The second principal component 

(13.17%) had high contributing factor loadings from 
Sep-ext-W. The third principal component account-
ed for 8.08% of the total variation.

The PCA ordination diagram of population sam-
ples (Fig. 2) indicated a slight differentiation of F. 
moschata and F. viridis populations along the first 
PC axis. The individuals of F. vesca are overlapping 
with individuals of F. moschata and F. viridis.

Canonical discriminant analyses have shown 
the separation of the three morphological groups of 
populations that match the analysed species (Fig. 2). 
Populations of F. viridis and F. moschata were dif-
ferentiated along the first DA axis, while the popula-

BIOLOGICA NYSSANA ● 10 (1) September 2019: 9-16 Nikolić et al. ● Morphometric analysis of vegetative and reproductive 
organs of the Fragaria species

14

Fig 2. Results of the principal component analysis (PCA) and canonical discriminant analysis (CDA) based on 
morphometric characters of the Fragaria species



BIOLOGICA NYSSANA ● 10 (1) September 2019: 9-16 Nikolić et al. ● Morphometric analysis of vegetative and reproductive 
organs of the Fragaria species

15

tion of F. viridis was slightly differentiated along the 
second axis. 

Cluster analysis of morphometric data has shown 
differentiation into two clusters, the first consisted of 
F. vesca and F. viridis, and the second included only 
F. moschata (Fig. 3).

Fig 3. Results of cluster analysis (UPGMA) for morpho-
logical characters of the Fragaria species based on Maha-
lanobis distances

Our study confirmed the presence of a high degree 
of variability in vegetative and reproductive charac-
ters of F. vesca populations, which is in agreement 
with the previous studies on this species (Sargent et 
al., 2004; Labokas & Bagdonaitë, 2005; Huseinović 
& Osmanović, 2010; Maliníková et al., 2013). The 
study of phenotypic diversity of F. vesca and F. 
viridis by Labokas & Bagdonaitë (2005) recognised 
some differences in the number of flowers, length of 
petioles, length and width of leaf between F. vesca 
and F. viridis. In our sample, inflorescences of F. 
vesca populations were composed of 6-12 flowers 
while F. viridis produced a lower number of flowers 
(4-7). In comparison to populations of F. viridis, F. 
vesca was characterised by shorter petioles, longer 
inflorescences and a higher ratio of peduncle to ro-
sette leaf length (Labokas & Bagdonaitë, 2005). Our 
study has shown different results regarding the range 
of variability of mentioned characters in F. vesca and 
F. viridis populations. Namely, the number of flow-
ers in F. vesca populations ranged from 1 to 7 and 
in F. viridis populations from 1 to 5. Also F. vesca 
individuals had longer petioles than F. viridis. Ac-
cording to the available literature, morphological 
variability of F. moschata was never studied before. 

This species has shown the highest level of variabil-
ity for almost all characters. F. moschata individuals 
had the highest values for characters pertaining to 
leaves (Leaf-L, Leaf-W, Lam-cent-L, Lam-cent-W, 
Lam-lat-L, Lam-lat-W) and almost all morphologi-
cal characters of the flowering part of the plant.

Conclusion
The morphological characters of vegetative and re-
productive organs of Fragaria species have shown 
high to moderate levels of variability. The most vari-
able morphological characters of vegetative organs 
were Peti-L and Leaf-L, while the most variable 
morphological characters of reproductive organs 
were Scap-L, Flow-branch-N, Flow-N, and Sep-
ext-Lob-N. Analysis of variance has shown that all 
morphometric characters except Lam-cent-lob-N 
contributed significantly to differentiation of ana-
lysed Fragaria species. Morphological characters 
responsible for differentiation of analyzed Fragaria 
species included: Leaf-L, Lam-cent-L, Lam-cent-W, 
Lam-lat-L, Ped-L and Sep-int-L.

Acknowledgements. This investigation is supported 
by the Ministry of Education, Science and Technologi-
cal Development of the Republic of Serbia grant number 
173030.

References

DiMeglio, L.M., Staudt, G., Yu, H., Davis, T.M. 
2014: A phylogenetic analysis of the genus Fra-
garia (strawberry) using intron-containing sequence 
from the ADH-1 gene. PLoS One. 9(7): e102237. 
Published 2014 Jul 31. doi:10.1371/journal.
pone.0102237
Gajić, M. 1972: Fragaria L. In: Josifović, M. (ed.), 
Flora SR Srbije, 4: 118-122, Srpska Akademija Nau-
ka i Umetnosti. Beograd.
Hancock, J.F., Bringhurst R.S. 1978: Inter-popula-
tional differentiation and adaptation in the perennial 
diploid species Fragaria vesca L. American Journal 
of Botany 65: 795-803.
Harrison, R.E., Luby, J.J., Furnier, G.R., Han-
cock, J.F. 1997: Morphological and molecular vari-
ation among populations of octoploid Fragaria vir-
giniana and Fragaria chiloensis (Rosaceaea) from 
North America. American Journal of Botany 84: 
612-620.
Harrison, R.E., Luby, J.J., Furnier, G.R., Han-
cock, J.F. 2000: Differences in the apportionment 
of molecular and morphological variation in North 
American strawberry and the consequences for ge-
netic resource management. Genetic Resources and 



16

BIOLOGICA NYSSANA ● 10 (1) September 2019: 9-16 Nikolić et al. ● Morphometric analysis of vegetative and reproductive 
organs of the Fragaria species

Crop Evolution 47: 647-657.
Huseinović, S., Osmanović, S. 2010: Morphomet-
ric and meristic characteristics of the Wild Strawber-
ry (Fragaria vesca L.) on Konjuh Mountain. Acta 
Agriculturae Serbica 15 (30): 133-140. 
Huseinović, S., Osmanović, S., Terzić, Z., 
Šabanović, M. 2014: Morphological and ecological 
differentiation of the fruit of Fragaria vesca L. (Ro-
saceae) from different habitats in Bosnia and Herze-
govina. Biologica Nyssana 5(2): 75-82.
Jenson R.J., Hancock, J.F. 1981: Multivariate rela-
tionships among California strawberries. Bulletin of 
the Torrey Botanical Club 109: 136-147.
Labokas, J., Bagdonaite, E. 2005: Phenotypic di-
versity of Fragaria vesca and F. viridis in Lithuania. 
Laboratory of Economic Botany, Institute of Botany 
3: 19-22. 
Maliníková, E., Kukla, J., Kuklová, M., Balážová 
M. 2013: Altitudinal variation of plant traits: mor-
phological characteristics in Fragaria vesca L. 
(Rosaceae). Annals of forest research 56(1): 79-89, 
2013.
Micevski K. 1998. Flora na Republika Makedonija, 
Tom I, sv. 4. Skopje: Makedonska Akademija na 

Naukite i Umetnostite.
Hummer, E.K., Bassil, N., Njuguna, W. 2011: Fra-
garia, In: Kole, C. (ed.), Wild Crop Relatives: Ge-
nomic and Breeding Resources, Temperate Fruits: 
17-44, Springer-Verlag Berlin Heidelberg.
Nosrati, H., Price, H.A., Gerstberger, P., Wilcock 
C.C. 2015: Characterization of an allotriploid straw-
berry Fragaria × bifera Duchesne (Rosaceae) from 
Europe. Natura Sloveniae 17(1):5-15.
Sargent, D.J., Geibel, M., Hawkins, J.A., Wilkin-
son, M.J., Battey, N.H., Simpson, D.W. 2004: 
Quantitative and qualitative differences in morpho-
logical traits revealed between diploid Fragaria spe-
cies. Annals of botany 94: 787-796.
Tutin, T.G., Heywood, V.H., Burges, N.A., Moore, 
D.M., Valentine, D.H., Walters, S.M., Webb, D.A. 
(eds.) 1964-1980: Flora Europaea, I-V. Cambridge 
University Press. London.
Zhang, Q., Folta, K.M., Davis, T.M. 2014: Somat-
ic embryogenesis, tetraploidy, and variant leaf mor-
phology in transgenic diploid strawberry (Fragaria 
vesca subspecies vesca ‘Hawaii 4’). BMC Plant Bi-
ology, 14: 1-11.