67
Key words: brambles, pollen 
aperture, exine ornamentation, 
light microscopy, scanning electron 
microscopy.
Ključne besede: robide, pelodna 
odprtina, ornamentacija eksine, 
svetlobna mikroskopija, vrstična 
elektronska mikroskopija.
Corresponding author:
Alla Hayrapetyan
E-mail:   
alla.hayrapetyan.63@gmail.com
Received: 26. 4. 2024
Accepted: 18. 11. 2024
Pollen morphological study of some 
species of the genus Rubus L. (Rosaceae) 
of the flora of Armenia
Abstract
Pollen morphology of 10 Armenian species of the genus Rubus L. was studied using 
both light microscopy (LM) and scanning electron microscopy (SEM), including 
the species R. candicans, R. cartalinicus and R. takhtadjanii investigated for the first 
time. The data received revealed the significant palynomorphological uniformity 
in the aperture type: mainly 3-zonocolporate, geniculate, occasionally 4-zonocolp-
orate one. In some 4-zonocolporate pollen grains of the species R. armeniacus, 
additional slit-like or colpus-like areas were also noted at one or both poles. Exine 
ornamentation was predominantly finely striate-perforate (SEM), sometimes in 
combination with finely striate-microreticulate one (R. armeniacus and R. caesius). 
Pollen grains were small, sometimes of medium size also. The smallest pollen grains 
(according to polar axis length) were observed in the species R. takhtadjanii (on 
average 14.2 μm), and the largest ones in R. caucasicus (on average 26.3 μm). Our 
data support the previously held view that pollen grain morphology within the 
genus Rubus can only be used as an auxiliary feature for the diagnosis of individual 
species.
Izvleček
S pomočjo svetlobne (LM) in vrstične elektronske mikroskopije (SEM) smo 
preučili morfologijo pelodnih zrn desetih vrst rodu Rubus iz Armenije. Pelod 
vrst R. candicans, R. cartalinicus in R. takhtadjanii smo preučili prvič. Dobljeni 
rezultati kažejo značilno palinomorfološko uniformnost tipa odprtine, večinoma 
3-zonokolporatne, genikulatne, občasno 4-zonokolporatne. Pri nekaterih 
4-kolporatnih pelodnih zrnih vrste R. armeniacus se pojavljajo še dodatne 
strukture, podobne brazdam na obeh polih. Ornamentacija eksine je bila večinoma 
striatno-perforatna (SEM), včasih v kombinaciji s fino striatno-mikroretikulatno 
(R. armeniacus in R. caesius). Pelodna zrna so bila majhna, včasih tudi srednje 
velika. Najmanjša pelodna zrna (glede na dolžino polarne osi) ima vrsta 
R. takhtadjanii (v povprečju 14,2 μm), vrsta R. caucasicus pa največja (v povprečju 
26,3 μm). Ugotovitve prejšnjih raziskav, da lahko morfologijo pelodnih zrn pri 
vrstah rodu Rubus uporabljamo samo kot pomožne znake za opis posameznih vrst, 
je potrdila tudi naša raziskava.
Alla Hayrapetyan1, Marieta Asatryan1, Hasmik Sonyan1, 
Karine Balayan1 & Dmitri Beketovski2
DOI: 10.3986/hacq-2025-000624/1 • 2025, 67–81
1 Institute of Botany after A. Takhtajyan, National Academy of Sciences of the Republic of Armenia, Yerevan, Republic of Armenia
2 Voskehat Educational and Research Center of Enology, Scientific branch of the Armenian National Agrarian University, Merdzavan, Republic of Armenia 
24/1 • 2025, 67–81
68
Hayrapetyan et al.
Pollen study of the genus Rubus L. in Armenia
Introduction
Rubus L. is one of the rather large and widespread genera 
of the Rosaceae family and is represented predominantly 
by erect to trailing shrubs or scramblers, rarely by herbs 
(Takhtajan, 2009). The genus is economically and eco-
logically important as fruit crops, ornamental plants, and 
as pioneers in early forest succession (Alice & Campbell, 
1999). It has also been revealed that Rubus readily invades 
natural areas (Daehler, 1998; Alice & Campbell, 1999). 
At the same time when exploring the pollen characteristics 
of the invasive in Poland anthropophyte species R. lacinia-
tus, Lechowicz et al. (2021) noted that pollen morphology 
probably does not affect the invasive traits of this species. 
Rubus species were a food source and medicinal plants for 
native peoples soon after the Ice Age (Connolly, 1999), 
and the medicinal use of brambles was documented in the 
writings of Aeschylus and Hippocrates, between 500 to 
370 BCE (Hendrickson, 1981; Hummer, 2010). It has 
been suggested that within subfamily Rosoideae Martinov 
the crown ages of its six tribes (including tribe Rubeae Du-
mort.) range from 45.02 to 10.42 Ma (the middle Eocene 
to the late Miocene) (Chen et al., 2020). According to Lu 
(1983, p. 25): “It is clearly shown that center of distribu-
tion lies in North America at present time”, from where 
the last common ancestor migrated to Central and South 
America, and Europe and Asia (Carter et al., 2019). 
According to various sources, the genus includes 250 to 
900 + species, with around 750 species that are document-
ed in Europe (Kurtto et al., 2011; Mabberley, 2017; Huang 
et al., 2023). Brambles exhibit a high level of morphologi-
cal diversity and are often characterized by hybridization, 
frequently associated with polyploidy and apomixis (Alice, 
2002; Sochor et al., 2015). The basic chromosome num-
ber is x = 7, and the genus has been reported to have dif-
ferent levels of ploidy in nature including 2x–18x as well 
as aneuploid genomes. Both natural and human-made 
hybrids are prevalent (Nybom, 1986; Foster et al., 2019; 
Gao et al., 2023). Specifically, “… except for a few sexual 
species occurring in Europe and its nearest vicinity… the 
subgenus Rubus is exclusively represented on the continent 
by polyploids of hybrid origin (2n = 21, 28, 35, and 42), 
where sexual reproduction has been almost entirely sup-
planted by apomixis” (Zieliński, 2004, p. 1). 
In some classification systems, Rubus is categorized un-
der the subfamily Ruboideae Thomé (Takhtajan, 1997, 
2009) and the tribe Rubeae (Angiosperm Phylogeny 
Website, https://www.mobot.org/mobot/research/ap-
web/), among others. The global taxonomic classification 
of the genus was established early in the 20th century, 
leading to the definition of 12 subgenera (Focke, 1910, 
1911, 1914). However, a recent phylogenetic study of 
145 Rubus species, including cultivars and hybrids, iden-
tified 10 subgenera (Huang et al., 2023). This study found 
that the genus is not monophyletic; only R. subgenus An-
oplobatus Focke forms a monophyletic group, while the 
remaining nine subgenera are either para- or polyphyletic 
(Chen et al., 2016; Huang et al., 2023). At the same time, 
according to Carter et al. (2019) R. subgenus Orobatus 
(Focke) Focke is also monophyletic. 
In Armenia, the total number of Rubus species is 13-14, 
the vast majority of which belong to the subgenus Ru-
bus (= Eubatus Focke) (Mulkijanyan, 1958), while spe-
cies R. idaeus L. belongs to the subgenus Idaeobatus Fo-
cke. Two rare endemic species, R. takhtadjanii Mulk. and 
R. zan gezurus Mulk., are listed in the Red Book of Plants 
of Armenia under the EN (endangered) category (Taman-
yan et al., 2010). For the species R. takhtadjanii, only one 
population is known in the Zangezur floristic region (in 
the vicinity of the villages Tsav and Srashen) on the terri-
tory of the Shikakhokh State Reserve and the Plane Grove 
Nature Reserve. The range of R. zangezurus is fragment-
ed, as the species grows both in the Ijevan floristic region 
(Berd) and in the Zangezur floristic region (Goris, Tsav) 
(Hayrapetyan et al., 2017). In the “Habitats of Armenia” 
(Fayvush & Aleksanyan, 2016) the genus is categorized 
under F2.33 (Subalpine mixed brushes), F2.338–AM 
(Subalpine crook stem forest), F3.2477–AM (Dewberry 
dense scrub), FB.3 (Shrub plantations for ornamental 
purposes or for fruit, other than vineyards), G1.371–AM 
(Plane grove in Tsav river valley), G1.927–AM (Aspen 
groves of North Armenia), G3.4E (Ponto–Caucasian 
Scots pine forests) and G5.85 (Shrubby  clearings). 
Pollen morphological features are often used to address 
complex taxonomical problems regarding interrelation-
ships between various taxa, particularly with reference 
to the families, subfamilies, tribes, genera, and species. 
The most distinctive characters include aperture type, ex-
ine ornamentation and the size of mature pollen grains. 
The earliest mention of the peculiarities of Rubus pollen 
morphology (using LM) can be found in the mid-20th 
century (Erdtman, 1952). Subsequently, numerous stud-
ies using a LM (e.g., Ikuse, 1955; Reitsma, 1966; Ku-
prianova, 1978) and a SEM (e.g., Hebda & Chinnappa, 
1990, 1994; Li et al. 2001; Wronska-Pilarek et al., 2012; 
Hanchana et al., 2023) have been conducted. In most 
cases, the authors highlight the size and shape of pollen 
grains as well as the exine ornamentation as the main di-
agnostic features. A large-scale study of pollen morphol-
ogy involving 155 species and 13 variations representing 
all 12 subgenera of Rubus was carried out by Xiong et 
al. (2019). The authors noted that pollen cluster analysis 
tree does not align well with the traditional macromor-
phological classification and molecular phylogenetic trees 
24/1 • 2025, 67–81
69
Hayrapetyan et al.
Pollen study of the genus Rubus L. in Armenia
based on DNA fragments. Other studies have similarly 
shown that, in general, pollen characters did not support 
the currently accepted taxonomic division of the genus 
Rubus into subgenera, sections and series (Lechowicz et 
al., 2022), as well as that “… pollen morphology alone is 
not sufficient to elucidate or reconstruct the taxonomic 
relationships within Rubus…” (Xiong et al., 2019, p. 
705). For that reason, pollen features in Rubus should be 
treated as auxiliary in taxonomy (Lechowicz et al., 2020). 
In Armenia, Avetisyan & Manukyan (1958) studied 
the pollen morphology of representatives of the genus 
Rubus using LM. The authors generally characterized the 
pollen of all investigated species as 3-zonocolporate with 
large pores and granulate exine ornamentation.
The main purpose of this study was to conduct a com-
parative palynomorphological analysis of 10 species of the 
genus Rubus in Armenia using LM and SEM to identify 
the main diagnostic characters that allow for the differen-
tiation of individual taxa.
Material and methods
Pollen morphological characteristics of 18 samples from 
10 species of the genus Rubus in Armenia were analyzed. 
Unopened mature flower buds were obtained from the 
herbarium of Institute of Botany after A. Takhtajyan, 
National Academy of Sciences of Republic of Armenia 
(ERE). 
The descriptions of pollen grains by light microscopy 
(LM) for each investigated species are based on acetolyzed 
material (Avetisyan, 1950) and also on grains stained with 
basic fuchsine (Smolyaninova & Golubkova, 1950) with 
obligatory fixation of pollen in glycerin jelly.
Six morphological characters, namely length of polar 
axis (P), length of equatorial diameter (E), length of col-
pi, apocolpium diameter, mesocolpium width and exine 
thickness, were measured.
Measurements under LM (AmScope 2000X LED, 
China, 2015) were taken on 20 pollen grains for each 
specimen. The studies were carried out at ×200, ×400 and 
×1000 magnifications. For descriptions of pollen shape, 
size, and aperture stained pollen grains were used; the 
details of exine structure and ornamentation were stud-
ied on acetolyzed pollen grains. For scanning electron 
microscopy (SEM), non-acetolyzed pollen grains were 
washed with alcohol, placed on a metal stub, and sputter 
coated with gold (10 nm). Samples were imaged under 
SEM (JEOL JSM-6390), with a 15 kV electron beam. 
Statistical analysis of all studied species was performed 
using Microsoft Excel with two indicators: ± SD – stan-
dard deviation and CV% – coefficient of variation. The 
sample is weakly variable in case of CV≤10%, with CV 
from 10% to 20% it is moderately variable, in case of 
CV≥20% it is being considered as highly variable and 
with CV≥30%, the highest degree of variability is noted.
For each of the species presented below, depending on 
the amount of available pollen material, one to four speci-
mens were studied. The list of investigated species and 
specimens is presented in Table 1; their names fully cor-
responding  to those presented in The Plant List (http://
www.theplantlist.org/). Data on some palynological fea-
tures of the genus Rubus in Armenia are presented in Ta-
ble 2; the average sizes of pollen grains (μm) of the stud-
ied species (P × E) using both our data and information 
from the literature are shown in Figure 1. The compari-
son of the average values for some pollen characteristics 
R.
 a
rm
en
ia
cu
s
R.
 a
rm
en
ia
cu
s (
3)
R.
 c
ae
siu
s
R.
 c
ae
siu
s (
1)
R.
 c
ae
siu
s (
2)
R.
 c
ae
siu
s (
4)
R.
 c
an
di
ca
ns
R.
 c
an
es
ce
ns
R.
 c
an
es
ce
ns
 (4
)
R.
 c
ar
ta
lin
ic
us
R.
 c
au
ca
sic
us
R.
 ib
er
ic
us
R.
 ib
er
ic
us
 (4
)
R.
 id
ae
us
R.
 id
ae
us
 (1
)
R.
 id
ae
us
 (2
)
R.
 sa
nc
tu
s
R.
 ta
kh
ta
dj
an
ii
40
30
20
10
0
Figure 1: Average sizes of pollen grains 
(P × E) in some species of the genus 
Rubus L.  
Our data: blue – polar axis (P), green 
– equatorial diameter (E). Data from 
literature: orange – polar axis (P), yellow 
– equatorial diameter (E). (1) – Li et al. 
(2001), (2) – Lechnovich et al. (2020), 
(3) – Tomlik-Wyremblewska (1995), 
(4) – Monasterio-Huelin & Pardo (1995).
Slika 1: Povprečna velikost pelodnih zrn 
(P × E) nekaterih vrst rodu Rubus L.  
Naši podatki: modra – polarna 
(vertikalna) os (P), zelena – ekvatorialna 
(horizontalna) os (E). Podatki iz 
literature: oranžna – polarna (vertikalna) 
os (P), rumena – ekvatorialna 
(horizontalna) os (E). (1) – Li et al. 
(2001), (2) – Lechnovich et al. (2020), 
(3) – Tomlik-Wyremblewska (1995), 
(4) – Monasterio-Huelin & Pardo (1995).
24/1 • 2025, 67–81
70
Hayrapetyan et al.
Pollen study of the genus Rubus L. in Armenia
of the species studied, namely, polar axis (P), equatorial 
diameter (E), apocolpium diameter and mesocolpium 
width, was carried out using Simpson and Roe test (Van 
der Pluym & Hideux, 1997) (Figures 2–5).
The classification of the shape and size of pollen grains 
was given according to Erdtman (1952). The morphologi-
cal terminology used in our study mainly follows Erdtman 
(1952), Kuprianova & Alyoshina (1967), Punt et al. (2007), 
Halbritter et al. (2018), and Hayrapetyan & Bruch (2020).
Karyological studies the genus Rubus in Armenia have 
not been conducted, and the chromosome numbers for 
some species, as shown in Table 2, have been obtained 
from literature sources. 
R.
 a
rm
en
ia
cu
s
R.
 c
ae
siu
s
R.
 c
an
di
ca
ns
R.
 c
an
es
ce
ns
R.
 c
ar
ta
lin
ic
us
R.
 c
au
ca
sic
us
R.
 ib
er
ic
us
R.
 id
ae
us
R.
 sa
nc
tu
s
R.
 ta
kh
ta
dj
an
ii
20
15
10
5
Figure 2: Comparison of the average values of pollen polar axis 
(P) of investigated Rubus species using Simpson and Roe test.
Slika 2: Primerjava povprečnih vrednosti polarne (vertikalne) osi 
(P) preučevanih vrst rodu Rubus s Simpsonovim in Roevim testom.
R.
 a
rm
en
ia
cu
s
R.
 c
ae
siu
s
R.
 c
an
di
ca
ns
R.
 c
an
es
ce
ns
R.
 c
ar
ta
lin
ic
us
R.
 c
au
ca
sic
us
R.
 ib
er
ic
us
R.
 id
ae
us
R.
 sa
nc
tu
s
R.
 ta
kh
ta
dj
an
ii
35
30
25
20
15
10
R.
 a
rm
en
ia
cu
s
R.
 c
ae
siu
s
R.
 c
an
di
ca
ns
R.
 c
an
es
ce
ns
R.
 c
ar
ta
lin
ic
us
R.
 c
au
ca
sic
us
R.
 ib
er
ic
us
R.
 id
ae
us
R.
 sa
nc
tu
s
R.
 ta
kh
ta
dj
an
ii
25
20
15
10
R.
 a
rm
en
ia
cu
s
R.
 c
ae
siu
s
R.
 c
an
di
ca
ns
R.
 c
an
es
ce
ns
R.
 c
ar
ta
lin
ic
us
R.
 c
au
ca
sic
us
R.
 ib
er
ic
us
R.
 id
ae
us
R.
 sa
nc
tu
s
R.
 ta
kh
ta
dj
an
ii
7
6
5
4
3
2
Species Herbarium sheet information
R. armeniacus Focke ERE, 80977; ERE, 93906
R. caesius L. ERE, 56383; ERE, 63234; ERE, 63238
R. candicans Weihe ex Rchb. ERE, 58307; ERE, 63240
R. canescens DC. ERE, 58846; ERE, 98402
R. cartalinicus Juz. ERE, 63221
R. caucasicus Focke ERE, 58313
R. ibericus Juz. ERE, 135970
R. idaeus L. ERE, 98394
R. sanctus Schreb. ERE, 56378; ERE, 56381; ERE, 63978; 
ERE, 98366
R. takhtadjanii Mulk. ERE, 64509
Table 1: List of investigated species and specimens of the genus 
Rubus (ERE – Herbarium of the Institute of Botany after 
A. Takhtajyan, National Academy of Sciences, Yerevan, Armenia).
Tabela 1: Seznam preučevanih vrst in primerkov rodu Rubus 
(ERE – Herbarij Botaničnega inštituta A. Takhtajyan, Nacion-
alna akademija znanosti, Erevan, Armenija).
Results
Pollen grains are 3-zonocolp-orate, sometimes 4-zono-
colp-orate (Figure 6b), often prolate or subprolate (the 
shape of pollen grains can vary even within the same 
sample), sometimes prolate spheroidal in R. cartalinicus 
and subspheroidal in R. takhtadjanii (Figure 1, Table 2); 
outline in polar view is rounded-3(4)-angular or round-
ed-3(4)-lobed; polar axis (P) varies from 12.9 μm (in R. 
takhtadjanii) to 33.5 μm (in R. sanctus) (Table 2, Figu-
re 2), equatorial diameter (E) from 12.1 μm (in R. san-
ctus) to 24.1 μm (in R. caesius) (Table 2, Figure 3). Colpi 
Figure 3: Comparison of the average values of 
pollen equatorial diameter (E) of investigated 
Rubus species using Simpson and Roe test.
Slika 3: Primerjava povprečnih vrednosti 
premera ekvatorialnega premera pelodnega zrna 
(E) preučevanih vrst rodu Rubus s Simpso-
novim in Roevim testom.
Figure 4: Comparison of the average values 
of pollen apocolpium diameter of investigated 
Rubus species using Simpson and Roe test.
Slika 4: Primerjava povprečnih vrednosti 
premera apokolpija (brazde) preučevanih vrst 
rodu Rubus s Simpsonovim in Roevim testom.
Figure 5: Comparison of the average values 
of pollen mesocolpium width of investigated 
Rubus species using Simpson and Roe test.
Slika 5: Primerjava povprečnih vrednosti 
premera mezokolpija (brazde) preučevanih vrst 
rodu Rubus s Simpsonovim in Roevim testom.
24/1 • 2025, 67–81
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Hayrapetyan et al.
Pollen study of the genus Rubus L. in Armenia
Sp
ec
ie
s
Po
la
r a
xi
s (
P)
 
(μ
m
)
± 
SD
 (C
V
%
)
Eq
ua
to
ria
l  
di
am
et
er
 (E
) 
(μ
m
)
± 
SD
 (C
V
%
)
P/
E
C
o
l
p
u
s
E
x
i
n
e
 
o
r
n
a
m
e
n
t
a
t
i
o
n
C
hr
om
os
om
e 
 
nu
m
be
r (
2n
)
ap
oc
ol
pi
um
 
 di
am
et
er
 (μ
m
) 
± 
SD
 (C
V
%
)
m
es
oc
ol
pi
um
 
w
id
th
 (μ
m
) 
± 
SD
 (C
V
%
)
LM
SE
M
R.
 a
rm
en
ia
cu
s 
Fo
ck
e
23
.5
–2
5.
8 
/ 2
4.
5 
± 
0.
7 
(2
.7
%
)
16
.9
–2
2.
8 
/ 1
9.
4 
±1
.8
 (9
.2
%
)
8/
7–
8/
6 
(s
ub
pr
ol
at
e)
4.
8–
5.
9 
/ 5
.3
 
± 
0.
4 
(7
.1
%
)
8.
4–
12
.5
 / 
10
.4
 
± 
1.
0 
(9
.8
%
)
fin
el
y 
re
tic
ul
at
e
fin
el
y 
str
ia
te
-p
er
fo
ra
te
 o
r 
fin
el
y 
str
ia
te
-m
ic
ro
re
tic
ul
at
e
28
 
(K
ra
hu
lc
ov
á 
&
 
H
ol
ub
, 1
99
7)
R.
 ca
esi
us
 L
.
22
.2
–2
8.
9 
/ 2
5.
1 
± 
1.
8 
(7
.2
%
)
19
.7
–2
4.
1 
/ 2
0.
9  
       
± 
1.
6 
(7
.4
%
)
8/
7–
8/
6 
(s
ub
pr
ol
at
e)
 
4.
5–
5.
3 
/ 4
.9
 
± 
0.
3 
(5
.3
%
)
14
.1
–1
5.
8 
/ 1
4.
9 
± 
0.
4 
(2
.8
%
)
fin
el
y 
re
tic
ul
at
e
fin
el
y 
str
ia
te
-p
er
fo
ra
te
 o
r 
fin
el
y 
str
ia
te
-m
ic
ro
re
tic
ul
at
e
28
 
(K
ra
hu
lc
ov
á 
&
 
H
ol
ub
, 1
99
7)
R.
 ca
nd
ica
ns
 
W
ei
he
 e
x 
R
ch
b.
20
.3
–2
5.
4 
/ 2
2.
9 
± 
1.
9 
(8
.4
%
)
17
.5
–2
2.
1 
/ 1
9.
0 
± 
1.
6 
(8
.7
%
)
8/
7–
8/
6 
(s
ub
pr
ol
at
e)
5.
0–
5.
6 
/ 5
.3
 
± 
0.
1 
(2
.8
%
)
14
.2
–1
5.
4 
/ 1
4.
8 
± 
0.
3 
(2
.0
%
)
gr
an
ul
at
e
fin
el
y 
str
ia
te
-p
er
fo
ra
te
21
 
(G
us
ta
fss
on
,1
94
3)
R.
 ca
ne
sce
ns
 D
C
.
23
.3
–2
5.
8 
/ 2
4.
5 
± 
0.
7 
(3
.0
%
)
17
.7
–2
0.
5 
/ 1
9 
± 
0.
9 
(4
.9
%
)
8/
7–
8/
6 
(s
ub
pr
ol
at
e)
 
4.
2–
5.
1 
/ 4
.7
 
± 
0.
2 
(4
.8
%
) 
(s
om
et
im
es
 
sy
na
pe
rt
ur
at
e)
10
.5
–1
4.
3 
/ 1
2.
4 
± 
1.
0 
(7
.7
%
)
fin
el
y 
str
ia
te
- 
re
tic
ul
at
e
fin
el
y 
str
ia
te
-p
er
fo
ra
te
, s
tr
ia
e 
lin
ke
d 
to
ge
th
er
 b
y 
br
id
ge
s
14
 
(K
ra
hu
lc
ov
á 
&
  
H
ol
ub
, 1
99
7)
R.
 ca
rt
al
in
icu
s 
Ju
z.
21
.8
–3
0.
1 
/ 2
5.
8 
± 
3.
1 
(1
1.
8%
)
22
.1
–2
3.
5 
/ 2
2.
9 
 
± 
0.
5 
(2
.0
%
)
8/
8–
8/
7 
(p
ro
la
te
 
sp
he
ro
id
al
)
2.
8–
4.
5 
/ 3
.6
 
± 
0.
4 
(1
1.
6%
) 
(s
om
et
im
es
 
sy
na
pe
rt
ur
at
e)
14
.4
–1
6.
8 
/ 1
5.
6 
± 
0.
6 
(3
.8
%
)
fin
el
y 
str
ia
te
- 
re
tic
ul
at
e
fin
el
y 
str
ia
te
-p
er
fo
ra
te
–
R.
 ca
uc
as
icu
s 
Fo
ck
e
21
.7
–3
3.
2 
/ 2
6.
3 
± 
4.
1 
(1
5.
7%
)
16
.8
–2
2.
5 
/ 1
8.
0 
± 
3.
6 
(1
9.
8%
)
8/
6–
8/
4 
(p
ro
la
te
)
4.
7–
5.
9 
/ 5
.2
4 
± 
0.
4 
(7
.5
%
)
18
.5
–1
9.
5 
/ 1
9.
0 
± 
0.
3 
(1
.7
%
)
gr
an
ul
at
e
fin
el
y 
str
ia
te
-p
er
fo
ra
te
–
R.
 ib
er
icu
s J
uz
.
20
.5
–2
2.
5 
/ 2
1.
4 
± 
0.
7 
(3
.1
%
)
14
.6
–1
8.
1 
/ 1
6.
2 
± 
1.
2 
(7
.6
%
)
8/
6–
8/
4 
(p
ro
la
te
)
3.
5–
4.
8 
/ 4
.2
 
± 
0.
3 
(7
.8
%
)
11
.8
–1
4.
6 
/ 1
3.
2 
± 
0.
7 
(5
.3
%
)
fin
el
y 
re
tic
ul
at
e
–
–
R.
 id
ae
us
 L
.
18
.5
–2
5.
3 
/ 2
1.
8 
± 
2.
5 
(1
1.
6%
)
14
.1
–1
8.
9 
/ 1
6.
5 
± 
1.
6 
(9
.6
%
)
8/
6–
8/
4 
(p
ro
la
te
)
3.
8–
5.
7 
/ 4
.8
 
± 
0.
5 
(1
0.
0%
)
10
.5
–1
4.
9 
/ 1
2.
7 
± 
1.
1 
(8
.7
%
)
fin
el
y 
re
tic
ul
at
e
–
14
, 2
1 
(C
he
n 
et
 a
l.,
 1
99
3;
  
Iv
an
ov
a,
 2
00
5)
R.
 sa
nc
tu
s 
Sc
hr
eb
.
15
.5
–3
3.
5 
/ 1
9.
2 
± 
5.
3 
(2
7.
7%
)
12
.1
–1
7.
5 
/ 1
4.
4 
± 
1.
6 
(1
1.
3%
)
8/
6–
8/
4 
(p
ro
la
te
)
4.
8–
6.
1 
/ 5
.4
 
± 
0.
3 
(6
.0
%
)
10
.9
–1
2.
5 
/ 1
1.
7 
± 
0.
4 
(3
.4
%
)
gr
an
ul
at
e
fin
el
y 
str
ia
te
-p
er
fo
ra
te
, 
di
am
et
er
 o
f p
er
fo
ra
tio
n 
va
rie
s
14
 
(R
un
em
ar
k,
 2
00
6)
R.
 ta
kh
ta
dj
an
ii 
M
ul
k.
12
.9
–1
5.
6 
/ 1
4.
2 
± 
0.
8 
(5
.6
%
)
14
.1
–1
6.
8 
/ 1
5.
4 
± 
0.
9 
(6
.1
%
)
6/
8–
8/
6 
(s
ub
sp
he
ro
id
al
)
5.
3–
6.
9 
/ 6
.1
 
± 
0.
4 
(6
.6
%
)
9.
8–
11
.5
 / 
10
.7
 
± 
0.
4 
(4
.0
%
)
fin
el
y 
re
tic
ul
at
e
–
–
Ta
bl
e 
2:
 S
om
e 
pa
ly
no
lo
gi
ca
l a
nd
 k
ar
yo
lo
gi
ca
l f
ea
tu
re
s o
f t
he
 re
pr
es
en
ta
tiv
es
 o
f t
he
 g
en
us
 R
ub
us
 L
. o
f t
he
 fl
or
a 
of
 A
rm
en
ia
 (±
 S
D
 –
 st
an
da
rd
 d
ev
ia
tio
n,
 C
V
%
 –
 c
oe
ffi
ci
en
t o
f 
va
ria
tio
n,
 L
M
 –
 li
gh
t m
ic
ro
sc
op
y, 
SE
M
 –
 sc
an
ni
ng
 e
le
ct
ro
n 
m
ic
ro
sc
op
y)
.
Ta
be
la
 2
: N
ek
at
er
e 
pa
lin
ol
oš
ke
 in
 k
ar
io
lo
šk
e 
zn
ač
iln
os
ti 
pr
ed
sta
vn
ik
ov
 ro
du
 R
ub
us
 L
. v
 fl
or
i A
rm
en
ije
 (±
 S
D
 –
 st
an
da
rd
ni
 o
dk
lo
n,
 C
V
%
 –
 k
oe
fic
ie
nt
 v
ar
ia
ci
je
, L
M
 –
 sv
et
lo
bn
i 
m
ik
ro
sk
op
, S
EM
 –
 v
rs
tič
ni
 e
le
kt
ro
ns
ki
 m
ik
ro
sk
op
).
24/1 • 2025, 67–81
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Figure 6: Pollen grains of Rubus armeniacus Focke. a–h – LM micrographs: a – 3-zonocolp-orate pollen grain, b – 4-zonocolp-orate pollen grain 
with additional small slit-like apertures at the pole (marked with arrows), c – spheroidal os (marked with arrow), d, e – thickened edges of colpi 
(marked with arrows), f – geniculum (marked with arrow), g – exine, columellae layer (marked with arrow), h – finely reticulate exine ornamen-
tation; i, j – SEM micrographs: i – pollen grain in equatorial view, colpus, j – striate-perforate exine ornamentation in combination with finely 
striate-microreticulate one; scale bars: a–i – 10 μm, j – 1 μm.
Slika 6: Pelodna zrna vrste Rubus armeniacus Focke. a–h – LM mikrografi: a – 3-zonokolporatna pelodna zrna, b – 4 zonokolporatna pelodna 
zrna z dodatno odprtino na polih (označeno s puščico), c – sferoidna os (označeno s puščico), d, e – odebeljen del brazde (označeno s puščico), 
f – genikulum (označeno s puščico), g – eksina, kolumelna plast (označeno s puščico), h – fino mrežasta ornamentacija eksine, i, j – SEM 
mikrograf: i – pelodno zrno v ekvatorialnem pogledu, brazda, j – črtasto-luknjičasta ornamentacija eksine v kombinaciji s fino črtasto-
mikroretikuliranim okrasjem; merilce: a–i – 10 μm, j – 1 μm.
are long or very long (colpus length 4/5–1 of polar axis), 
occasionally anastomose towards the poles (i.e., synaper-
turate) (Figure 9b; Figure 10b, c), often geniculate (i.e., 
with bulge in the equatorial exine of the colpus) (Fig-
ure 6f; Figure 8e, g; Figure 9d), from rather narrow to 
narrow, sometimes almost slit-like (rarely wide), usually 
with evenly thickened edges (Figure 6d, e; Figure 9e; 
Figure 10e) and with rounded or pointed ends; colpus 
membrane ornamentation from psilate to irregularly ver-
rucate (Figure 10a); apocolpium diameter from 2.8 μm 
(in R. cartalinicus) to 6.9 μm (in R. takhtadjanii) (Ta-
ble 2, Figure 4), mesocolpium width from 8.4 μm (in 
R. armeniacus) to 19.5 μm (in R. caucasicus) (Table 2, 
Figure 5). Ora are spheroidal or oblate spheroidal (Fig-
ure 7c; Figure 9c), due to the presence of the geniculum 
or the convergence of the colpi at the equator, sometimes 
weakly expressed, rarely with uneven edges and indis-
tinct ends. Exine thickness is 1.2–1.3 μm, columellae 
separate, with spherical or claviform heads (Figure 6g; 
Figure 8c). Exine ornamentation is granulate (Figure 7f), 
reticulate (Figure 6h) or striate-reticulate (Figure 9f) 
(LM); in the vast majority of the species ornamentation is 
finely striate-perforate (Figure 8h; Figure 9h; Figure 10g; 
Figure 11e), sometimes (R. armeniacus and R. caesius) in 
combination with finely striate-microreticulate one (Fig-
ure 6j; Figure 7h) (SEM).
 The striae are long, often dichotomously branched; in 
the species R. canescens, the individual striae are connect-
ed by thin transverse bridges (Figure 9h). The diameter 
of the perforations is 0.1–0.7 μm; the strongest variabil-
ity of this character was found in the species R. sanctus 
(Figure 11e). 
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Figure 7: Pollen grains of Rubus caesius L. a–f – LM micrographs: a–b – pollen grains in polar view (a – colpi, b – exine, columellae layer), 
c – spheroidal os (marked with arrow), d–f – pollen grains in equatorial view (e – os area, marked with arrows), f – granulate exine ornamenta-
tion; g–h – SEM micrographs: g – pollen grains in polar and equatorial view, h – striate-perforate exine ornamentation in combination with finely 
striate-microreticulate one; scale bars: a–g – 10 μm, h – 1 μm.
Slika 7: Pelodna zrna vrste Rubus caesius L. a–f – LM mikrografi: a–b – pelodno zrno v polarnem pogledu (a – brazda, b – eksina, kolumelna plast), 
c – sferoidna os (označena s puščico), d–f – pelodno zrno v ekvatorijalnem pogledu (e – osno območje, označeno s puščico), f – zrnata ornamentaci-
ja eksine, g–h – SEM mikrograf: g – pelodno zrno v polarnem in ekvatorialnem pogledu, h – črtasto-luknjičasta ornamentacija eksine v kombinaciji 
s fino črtasto-mikroretikulirano ornamentacijo; merilce: a–g – 10 μm, h – 1 μm.
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Pollen study of the genus Rubus L. in Armenia
Figure 8: Pollen grains of Rubus candicans Weihe ex Rchb. a–f – LM micrographs: a–c – pollen grains in polar view (a – colpus membrane 
 ornamentation, b – os area, c – exine, columellae layer, marked with arrows), d–f – pollen grains in equatorial view (d – colpus without genuculum, 
e – colpus with genuculum, marked with arrows, f – thickened edges of colpi); g–h – SEM micrographs: g – pollen grains in polar and equatorial 
view (colpus with genuculum, marked with arrow), h – finely striate-perforate exine ornamentation (SEM); scale bars: a–g – 10 μm, h – 1 μm.
Slika 8: Pelodna zrna vrste Rubus candicans Weihe ex Rchb. a–f – LM mikrografi: a–c – pelodna zrna v polarnem pogledu (a – ornamentacija 
 membrane brazde, b – območje osi, c – eksina, kolumelna plast, označeno s puščico), d–f – pelodno zrno v ekvatorialnem pogledu (d – brazda brez 
genikuluma, e – brazda z genikulumom, označeno s puščico, f – odebeljen del brazde);  g–h – SEM mikrograf: g – pelodna zrna v polarnem in 
ekvatorialnem pogledu (brazda z genikulumom, označeno s puščico), h – fino črtasto-luknjasta ornamentacija eksine (SEM); merilce: a–g – 10μm, 
h – 1 μm.
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Figure 9: Pollen grains of Rubus canescens DC. a–f – LM micrographs: a – pollen grains in semipolar view, b – synaperturate pollen grain in polar 
view (synaperture marked with arrow), c–f – pollen grains in equatorial view (c – os, d – genuculum, e – thickened edges of colpi, marked with 
 arrows, f – striate-reticulate exine ornamentation); g–h – SEM micrographs: g – pollen grains in semipolar and equatorial view, h – finely striate-
perforate exine ornamentation (SEM); scale bars: a–g – 10 μm, h – 1 μm.
Slika 9: Pelodna zrna vrste Rubus canescens DC. a–f – LM mikrografi: a – pelodno zrno v semipolarnem pogledu, b – po daljšnici anastamozna 
 odprtina (označeno s puščico), c–f – pelodno zrno v ekvatorialnem pogledu (c – os, d – genikulum, e – odebeljeni rob brazde, označen s puščico, f – 
črtasto-mrežasta ornamentacija eksine; g–h SEM mikrograf: g – pelodno zrno v semipolarnem in ekvatorialnem pogledu, h – fino črtasto-luknjasta 
ornamentacija eksine (SEM); scale bars: a–g – 10 μm, h – 1 μm.
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Figure 10: Pollen grains of Rubus cartalinicus Juz. a–e – LM micrographs: a–c – pollen grains in polar view (a – pollen and colpus membrane 
ornamentation, b–c – synaperturate pollen grains and os area, marked with arrows), d–e – pollen grains in equatorial view (d – os, e – thickened 
edges of colpi); f–g – SEM micrographs: f – pollen grains in equatorial and semipolar view, g – finely striate-perforate exine ornamentation; scale 
bars: a–f – 10 μm, g – 1 μm.
Slika 10: Pelodna zrna vrste Rubus cartalinicus Juz. a–e – LM mikrografi: a–c – pelodno zrno v polarnem pogledu (a – pelod in ornamentacija 
membrane brazde, b–c pelodno zrno z anastamozno odprtino, označeno s puščico), d–e – pelodno zrno v ekvatorialnem pogledu (d – os, 
e – odebeljen del brazde); f–g – SEM mikrograf: f – pelodno zrno v ekvatorialnem in semipolarnem pogledu, g – fino črtasto-luknjasta 
ornamentacija eksine; merilce: a–f – 10 μm, g – 1 μm.
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Discussion
Apertures. Our studies revealed the presence of predomi-
nantly 3(4)-zonocolporate pollen apertures in representa-
tives of the genus Rubus from the Armenian flora, where 
endoapertures are formed due to thinning or complete 
absence of the internal layers of the exine. Addition-
ally, in some 4-zonocolporate pollen grains of the spe-
cies R.  armeniacus, additional slit-like or colpus-like areas 
were observed at one or both poles (Figure 6b), indicating 
a possible hybridogenic nature of this species. It should 
also be noted that such spheroidal or oblate spheroidal 
endoapertures, often invisible beyond the colpi, have been 
considered by several authors as pores (e.g., Avetisyan & 
Manukyan, 1958; Kuprianova, 1978; Kosenko et al., 
1984; Tomlik-Wyremblewska, 1995). On the other hand, 
pollen studies of five West-European Rubus species (Reits-
ma, 1966), as well 18 species from the Iberian Peninsula 
(Monasterio-Huelin & Pardo, 1995) revealed lalongate 
endoapertures formed due to the disappearance or thin-
ning of the nexine, thereby confirming our standpoint. 
Some authors characterized these areas as endopores with 
irregular margins and a fastigium (Wronska-Pilarek et al., 
2006; Wronska-Pilarek et al., 2012). 
A characteristic feature of pollen grains of the genus 
Rubus is the presence of a geniculum, i.e., a bulge in the 
equatorial exine of the colpus, usually formed due to the 
separation of the sexine from the nexine and the ruptu-
re of the latter (Potonié, 1934; Punt et al., 2007). The 
presence of a similar bulge in some species of the genus 
Rubus, referred to as “fastigium”, was noted by Reitsma 
for the first time (1966). It is possible that the presence 
of the geniculum covering the ora led several authors to 
characterize Rubus pollen as tricolpate (e.g., Demchenko, 
1967; Eide, 1981b). Some authors noted the presence 
of an equatorial bridge, with margins constricted at the 
equator (Tomlik-Wyremblewska, 1995).
Shape and size. The general outline of pollen grains in 
the investigated species was mainly prolate (P/E = 8/6–
8/4) or subprolate (P/E = 8/7–8/6), prolate spheroidal 
Figure 11: Pollen grains of Rubus sanctus Schreb. a–c – LM micrographs: a – pollen grain in polar view, b–c – pollen grains in equatorial view 
(c– colpus with genuculum, marked with arrow); d–e – SEM micrographs: d – pollen grain in equatorial view, e – finely striate-perforate exine 
ornamentation; scale bars: a–d – 10 μm, e – 1 μm.
Slika 11: Pelodna zrna vrste Rubus sanctus Schreb. a–c – LM mikrografi: a – pelodno zrno v polarnem pogledu, b–c – pelodno zrno v ekvatorial-
nem pogledu (c– brazda z genikulumom, označeno s puščico); d–e – SEM mikrograf: d – pelodno zrno v ekvatorialnem pogledu, e – fino črtasto-
luknjasta ornamentacija eksine; merilce: a–d – 10 μm, e – 1 μm.
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Pollen study of the genus Rubus L. in Armenia
(P/E = 8/8–8/7) in R. cartalinicus, and subspheroidal 
(6/8–8/6) in R. takhtadjanii (Table 2, Figure 1). 
In most cases, pollen grains were small. However, for 
R. caesius, R. cartalinicus Juz., R. caucasicus Focke, R. 
sanctus both small and medium-sized pollen were noted, 
with Pmax= 33.5 μm observed in R. sanctus (Table 2). The 
smallest pollen grains (according to polar axis length) 
were noted in the rare endemic species R. takhtadjanii 
(on average 14.2 μm), and the largest ones in R. cauca-
sicus (on average 26.3 μm). According to available liter-
ature data, the largest pollen grains in the genus Rubus 
(P × E = 42.5 × 35.8 μm) were found in the species R. 
alceifolius Poir. (Kosenko et al., 1984). 
 In general, according to Monasterio-Huelin & Pardo 
(1995, p. 231), “the comparative analysis of the mean val-
ues of P and E is not significant in distinguishing pollen 
types”.
Exine. The thickness of the exine layer in the species 
studied ranged from 1.2 to 1.3 μm, resulting in a certain 
percentage of destroyed pollen grains during acetolysis 
treatment According to literary sources, the maximum 
exine thickness was noted in the species R. nutkanus Moc. 
(2.5 μm) (Demchenko, 1967).
Exine ornamentation. Examination of the surface of 
pollen grains using SEM revealed finely striate-perforate 
exine ornamentation, sometimes (in R. armeniacus, R. 
caesius) combined with finely striate-microreticulate one 
(Figure 6j, Figure 7h). A similar type of sculpture is also 
noted for the pollen of approximately twenty species of 
the genus Rubus in the Iberian Peninsula (Monasterio-
Huelin & Pardo, 1995). The exception is R. genevierii 
Boreau, “… whose ornamentation is striate-gemmate-
perforate and with a striated margin” (Monasterio-
Huelin & Pardo, 1995, p. 229). Pollen studies of Rubus 
representatives have also revealed some other different 
types of ornamentation besides the perforate-stiate one, 
including rugulate, rugulate-striate, striate-scabrate, 
stiate-gemmate, stiate-reticulate, rugulate, reticulate, 
clavate-bacculate (Gonzalez Romano & Candau, 1989; 
Ueda, 1992; Tomlik-Wyremblewska, 1995, 2000, 2004; 
Zhou et al., 1999; etc.).
A wide-ranging research of the pollen morphology of 
155 species and 13 varieties representing all 12 subgen-
era of Rubus revealed six types of exine ornamentation 
(I – perforate, II – microreticulate, III – striate-perforate, 
IV – rugulate-perforate, V – echinate, VI – striate-micro-
reticulate) and three subtypes (IA– scabrate-perforate, IB 
– perforations and discontinuous muri; IIIA – scabrate-
striate-perforate) (Xiong et al., 2019). 
It is possible that the pronounced polyploidy has played 
an undoubted role in the display of pollen polymorphism 
within the genus Rubus. For example, in contrast to the 
striate or reticulate sculpture, sometimes a vermiculate 
sculpture is characteristic of the pollen of many species 
(Ghosh & Saha, 2017; Valdés et al., 1987; Wronska-Pi-
larek et al., 2012); the systematically and ecologically iso-
lated dioecious octoploid glacial relict R. chamaemorus L. 
(2n = 8x = 56) (Thiem, 2003) exhibits spinose (Erdtman, 
1952; Demchenko, 1967; Faegri & Iversen, 1989), ver-
rucate, gemmate (Reitsma, 1966; Hebda & Chinnappa, 
1990, 1994) or perforate-striate (Eide, 1981a) pollen ex-
ine ornamentations.
Disagreements also arise when characterizing the type 
of apertures in the aforementioned species, namely, 
3-colpor(oid)ate (Erdtman, 1952) vs tricolpate (Dem-
chenko, 1967). The latter author also noted clear differ-
ences in pollen characteristics between R. chamaemorus 
and two closely related species, R. trifidus Thunb. and R. 
nutkanus (synonym of R. parviflorus Nutt.), both in terms 
of aperture type (tricolpate vs tricolp-orate) and exine 
ornamentation type (spinose vs reticulate-foveolate and 
sinuously reticulate, respectively). Additionally, tricolpate 
pollen grains (Eide, 1981b) and tuberculate sculpture for 
the species R. arcticus L. (Surova, 1975), as well as micro-
verrucate exine ornamentation for R. parviflorus (Hebda 
& Chinnappa, 1990) were reported. In both R. idaeus 
(Eide, 1981b) and R. sanctus (our studies), a significant 
diversity in the total size of pollen grains within the same 
sample (almost twice) was observed. A considerable vari-
ability in pollen grain sizes between the different samples 
of species R. caesius and R. idaeus is shown in Figure 1. 
On the other hand, a study of pollen morphology and 
pollen variability in R. gracilis from 13 natural Polish lo-
calities “… revealed no differences among the grains from 
the individual localities, likely the result of apomixis” 
(Wronska-Pilarek et al., 2006, p. 69). 
Conclusions
Pollen morphology of 10 Armenian species of the genus 
Rubus was studied using both light microscopy (LM) and 
scanning electron microscopy (SEM), including the spe-
cies R. candicans, R. cartalinicus and R. takhtadjanii inves-
tigated for the first time. 
Analysis of the data obtained revealed significant paly-
nomorphological uniformity both in the type of aper-
tures (mainly 3(4)-zonocolporate) and the type of exine 
ornamentation (finely striate-perforate), in combination 
with finely striate-microreticulate one in R. armeniacus 
and R. caesius. On the other hand, the obtained data con-
firmed the observations of Wronska-Pilarek et al. (2012) 
and Xiong at al. (2019) regarding the significant variabil-
ity of certain pollen features within individual species of 
the genus Rubus (such as pollen shape, size, etc.), and es-
24/1 • 2025, 67–81
79
Hayrapetyan et al.
Pollen study of the genus Rubus L. in Armenia
pecially within the subgenus Rubus (= Eubatus), to which 
most of the Armenian species of the genus Rubus belong. 
For example, in R. sanctus a significant variability in the 
length of the polar axis (P) was revealed, ranging from 
15.5 to 33.5 μm.
In this context, our data support the previously held 
view that pollen grain morphology within the genus Ru-
bus can only be used as an auxiliary feature for the diag-
nosis of individual species.
Acknowledgements
We thank Sh. Ohanyan for technical assistance with LM 
photography and L. Kartzeva for assistance with SEM 
photography. We are also grateful to the anonymous ref-
erees for their suggestions, which helped to substantially 
improve the original manuscript. 
We would like to thank Nina Caf for the translation 
into Slovenian.
ORCID iDs
Alla Hayrapetyan  https://orcid.org/0000-0002-8038-7422
Marieta Asatryan  https://orcid.org/0009-0009-3875-3677
Hasmik Sonyan  https://orcid.org/0009-0007-9442-3424
Karine Balayan  https://orcid.org/0009-0002-7875-1085
Dmitri Beketovski  https://orcid.org/0009-0007-1660-8204
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