Мікроморфологічні особливості поверхні листка видів роду Sansevieriа Thunb. s.str. (Asparagaceae)
This study investigated the leaf surface micromorphology of 12 species of the genus Sansevieria using light microscopy and scanning electron microscopy. The research focused on identifying micromorphological traits associated with plant stress tolerance, including epidermal cell shape, cuticle thick...
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M.M. Gryshko National Botanical Garden of the NAS of Ukraine
2026
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| author | Maryniuk, Myroslava Buyun, Lyudmyla Gyrenko, Oleksandr Gurnenko, Ivan |
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| description | This study investigated the leaf surface micromorphology of 12 species of the genus Sansevieria using light microscopy and scanning electron microscopy. The research focused on identifying micromorphological traits associated with plant stress tolerance, including epidermal cell shape, cuticle thickness, stomatal distribution and density, and cuticular characteristics with epicuticular wax deposits. In most of the studied Sansevieria species, the leaves are amphistomatic, whereas hypostomatic leaves are observed in S. cylindrica, S. canaliculata, and S. suffruticosa. In all studied taxa, the epidermis consists of a single layer of cells and lacks trichomes.The examined Sansevieria species are characterized by a well-developed cuticular layer and the presence of wax deposits that perform protective and water-conserving functions. The thickness of the cuticle and its ornamentation vary both among species and between leaf surfaces within the same species. The abaxial leaf surface generally exhibits a more developed cuticle than the adaxial surface, a feature particularly pronounced in S. cylindrica, S. canaliculata, S. kirkii, S. roxburghiana, S. gracilis, S. suffruticosa, and S. intermedia. All investigated species possess anomocytic stomata. Stomatal density on the abaxial leaf surface ranged from 9 to 27 mm2 among the studied species. These interspecific variations reflect distinct strategies for optimizing water balance under arid conditions.At the level of leaf micromorphology, amphistomaty, the spatial organization of epidermal cells, the presence of a cuticular layer with epicuticular wax deposits of various configurations, differences in stomatal sunkenness and density, and the occurrence of underdeveloped stomata can be considered markers of stress tolerance in this genus. The identified micromorphological markers provide insight into the adaptive xeromorphic traits of Sansevieria and have potential applications in applied research, including biotechnological projects and phytoremediation, including green infrastructure development. |
| doi_str_mv | 10.46341/PI2025024 |
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© The Authors. This content is provided under CC BY 4.0 license.
Plant Introduction, 109, 20–34 (2026) ISSN 1605-6574, e-ISSN 2663-290X
RESEARCH ARTICLE
Micromorphological traits of the leaf surface in species of the genus
Sansevieria Thunb. s.str. (Asparagaceae)
Myroslava Maryniuk *, Lyudmyla Buyun, Oleksandr Gyrenko, Ivan Gurnenko
M.M. Gryshko National Botanical Garden of the NAS of Ukraine, Sadovo-Botanichna str. 1, 01103 Kyiv, Ukraine;
* maryniukmyroslava@gmail.com
Received: 19.12.2025 | Accepted: 16.01.2026 | Published: 18.01.2026
Abstract
This study investigated the leaf surface micromorphology of 12 species of the genus Sansevieria using light
microscopy and scanning electron microscopy. The research focused on identifying micromorphological
traits associated with plant stress tolerance, including epidermal cell shape, cuticle thickness, stomatal
distribution and density, and cuticular characteristics with epicuticular wax deposits. In most of the
studied Sansevieria species, the leaves are amphistomatic, whereas hypostomatic leaves are observed in
S. cylindrica, S. canaliculata, and S. suffruticosa. In all studied taxa, the epidermis consists of a single layer of
cells and lacks trichomes.
The examined Sansevieria species are characterized by a well-developed cuticular layer and the presence of
wax deposits that perform protective and water-conserving functions. The thickness of the cuticle and its
ornamentation vary both among species and between leaf surfaces within the same species. The abaxial
leaf surface generally exhibits a more developed cuticle than the adaxial surface, a feature particularly
pronounced in S. cylindrica, S. canaliculata, S. kirkii, S. roxburghiana, S. gracilis, S. suffruticosa, and S. intermedia.
All investigated species possess anomocytic stomata. Stomatal density on the abaxial leaf surface ranged
from 9 to 27 mm2 among the studied species. These interspecific variations reflect distinct strategies for
optimizing water balance under arid conditions.
At the level of leaf micromorphology, amphistomaty, the spatial organization of epidermal cells, the
presence of a cuticular layer with epicuticular wax deposits of various configurations, differences in
stomatal sunkenness and density, and the occurrence of underdeveloped stomata can be considered
markers of stress tolerance in this genus. The identified micromorphological markers provide insight
into the adaptive xeromorphic traits of Sansevieria and have potential applications in applied research,
including biotechnological projects and phytoremediation, including green infrastructure development.
Keywords: Sansevieria, light microscopy, scanning electron microscopy, cuticle, epidermis, stomatal apparatus, micromorphology,
phytoremediation
https://doi.org/10.46341/PI2025024
UDC 582.573.36 : 581.821
Authors’ contributions: Myroslava Maryniuk and Lyudmyla Buyun developed the concept and structure of this study, interpreted the
results, and prepared the draft manuscript. Myroslava Maryniuk and Ivan Gurnenko conducted the study using light and scanning
electron microscopy and ensured the visualization of the results. Oleksandr Gyrenko performed statistical processing of SEM results
using ImageJ software and participated in the preparation of illustrations. All authors of the publication participated in the discussion
of the manuscript.
Funding: The research was conducted as part of the govermental research project No. 409-OP “Induction of phenotypic plasticity
in tropical and subtropical plants: search for predictive markers of resistance” 2023–2027, state registration number 0123U101383.
Competing Interests: The authors declare no conflict of interest.
https://creativecommons.org/licenses/by/4.0/
https://orcid.org/0000-0003-2590-448X
https://orcid.org/0000-0002-9158-6451
https://orcid.org/0000-0003-3296-3787
https://orcid.org/0000-0001-7600-6909
Plant Introduction • 109 21
Micromorphological traits of the leaf surface in Sansevieria species
Introduction
Representatives of the genus Sansevieria s.str.
have been actively studied in recent years
with a view to their use in phytoremediation
(Boraphech & Thiravetyan, 2015; Tariq et al.,
2017), phytochemistry and phytopharmacology
(Tkachenko et al., 2017; Maryniuk et al.,
2018; Thu et al., 2020), biotechnology, and
ornamental horticulture. To date, the general
morphology of representatives of the genus
(Koller & Rost, 1988a), their general anatomy
(Koller & Rost, 1988b; Klimko et al., 2018) and
the structure of water-storing tissue in leaves
(Koller & Rost, 1988b), as well as the structure
of the epidermis of individual species
(Shkrum et al., 2012; Klimko et al., 2018). The
vast majority of works are devoted to the
cultivation of S. trifasciata Prain varieties
(Babu & Srinivasa Prabhu, 2023).
However, despite the large number of
publications devoted to various aspects of
the biology of this genus, modern anatomical
and morphological studies of the genus
Sansevieria remain insufficient, especially in
the context of searching for functional traits
that could be used in plant introduction for the
purpose of ex situ biodiversity conservation,
phytoremediation (thanks to ability of these
plants to withstand abiotic stresses and
absorb heavy metals), pharmacognosy, and
biotechnological developments as donors
of stable traits for in vitro cultivation or the
search for biologically active substances.
The genus Sansevieria s.str. comprises ca.
100 species and infraspecific taxa, most of
which have been described over the last three
decades (Newton, 2020; Burkart et al., 2023).
According to recent molecular phylogenetic
studies, the genus Sansevieria Petagna was
nested within the genus Dracaena Vand. ex L.
(Takawira Nyenya et al., 2018; van Kleinwee
et al., 2022), which belongs to the family
Asparagaceae, subfamily Nolinoideae, order
Asparagales (Kim et al., 2010; Chen et al.,
2013; Lu & Morden, 2014; The Angiosperm
Phylogeny Group et al., 2016). Within this
work, we use the term Sansevieria to refer
to the monophyletic group within Dracaena
– the Sansevieria clade (van Kleinwee
et al., 2022).
Sansevieria species occupy a wide
range of ecological niches. This is a group
of xerophytic, perennial succulent plants,
distributed mainly in tropical and subtropical
Africa, Madagascar, the Arabian Peninsula, and
the Indian subcontinent. Most representatives
of the genus are adapted to conditions of
limited water supply. Typical habitats of
Sansevieria species include termite mounds,
river floodplains, and rocky outcrops. Some
species form clumps at the base of tree trunks
(Takawira & Nordal, 2002). Members of the
Sansevieria genus are found in various tropical
biomes within so-called ‘hotspots’ that serve
as priority areas for biodiversity conservation
(Burkart et al., 2025).
Leaf micromorphology is one of the
key indicators of plant adaptation to
environmental conditions. During evolution,
certain structural features have developed
that correlate well with indicators of plant
water status, drought resistance, the ability
to regulate gas exchange, and the ability to
withstand stress factors.
Data on the micromorphological features of
the leaf were used to differentiate species of
the genus Phaedranassa Herb. (Amaryllidaceae,
Asparagales) along an altitude gradient (Zhila
& Marinyuk, 2017), to assess the ‘adaptation
syndrome’ when transferring juvenile plants
of Guarianthe bowringiana (O’Brien) Dressler
& W.E.Higgins (Orchidaceae) from in vitro
culture to greenhouse conditions (Buyun
et al., 2021). The taxonomic significance of the
micromorphological features of the vegetative
organs of Dracaena s.l. (Klimko et al., 2018),
as well as other genera of the Asparagaceae
family, including Aspidistra Ker Gawl.
(Vislobokov et al., 2021) and Chlorophytum
Ker Gawl. (Nalawade & Gurav, 2017) has been
proved earlier.
Therefore, our work aimed to conduct a
comparative study of the micromorphological
features of the leaf blade of Sansevieria
model species as a marker of adaptive traits
for use in biodiversity conservation and eco-
biotechnology.
Material and methods
The study materials were collected in the
greenhouse of the Department of Tropical
and Subtropical Plants at the M.M. Gryshko
National Botanical Garden of the National
Academy of Sciences of Ukraine. The objects
of the study were plants of 12 Sansevieria
22 Plant Introduction • 109
Maryniuk et al.
species: S. aethiopica Thunb., S. canaliculata
Thunb., S. cylindrica Bojer ex Hook., S. gracilis
N.E.Br., S. grandis Hook.f., S. intermedia N.E.Br.,
S. kirkii Baker, S. parva N.E.Br., S. roxburghiana
Schult. & Schult.f., S. suffruticosa N.E.Br.,
S. trifasciata Thunb., S. metallica Gérôme &
Labroy.
Mature leaves of the middle formation were
used for the study. The samples were taken
from the central part of the leaf blade.
When studying the leaf blade surface with
light microscopy (LM), the Palacci replica
technique was used, in which the collodion
solution was replaced with a transparent
varnish (Furst, 1979; Karupu, 1984). The scraping
method described by Cutler et al. (2007) was
also applied with minor modifications for
sample preparation.
For scanning electron microscopy (SEM)
examination, the samples were immersed
in 100 % tert-butanol for 1 h at –10 °C. Metal
containers with frozen samples were placed
on a table cooled to –60 °C in the working
volume of a vacuum universal post (VUP-5M).
During vacuuming of the working volume, the
samples were dried by sublimation of frozen
tert-butanol. The dried leaf samples were
coated with carbon in VUP-5M and platinum
using a JFC-1600 Auto Fine Coater (JEOL,
Japan). The micromorphology of the leaf
surfaces of the studied Sansevieria species
was visualized using a JSM-6700F scanning
electron microscope (JEOL, Japan) at an
acceleration voltage of 15 kV in high vacuum
mode.
The epidermal surface of the leaves of the
study species was described using terminology
commonly accepted in plant cytology (Fahn,
2002; Dickison, 2000). The density of stomata
and their linear dimensions were determined
using ImageJ software. The types of stomata
were determined according to Baranova’s
(1985) classification. Epicuticular wax deposits
were evaluated by applying the Barthlott et al.
(1998) technique.
For biometric studies of epidermal cells and
the stomatal apparatus, a Primo Star B 48-
0071 (Carl Zeiss) light microscope equipped
with a Canon PowerShot A640 digital camera
was used. Measurements were performed
using licensed AxioVision Rel. 4.7 (Carl Zeiss)
software.
The stomatal index (SI) was determined
using the Willmer (1983) formula:
SI = S/(E+S) × 100 %, where
SI – stomatal index;
S – number of stomata per unit of leaf
surface area;
E – number of ordinary epidermal cells,
including secondary cells, per unit of area.
Statistical processing of the data was
performed using standard methods with Excel
2010 software.
Results
Epidermis and ordinary epidermal cells
The epidermis of all studied taxa is
represented by a single layer of cells without
trichomes. The ordinary epidermal cells on
the adaxial and abaxial surfaces are oriented
along the long axis of the leaf and are arranged
in more or less regular rows (Figs. 1 & 2). The
cells are polygonal (4–7 angles), varying in
elongation, with anticlinal walls ranging from
straight to slightly curved. In most species,
the anticlinal walls are thickened, except in
S. parva, S. trifasciata, and S. metallica (Fig. 2
A–D, H, I).
The dimensions of epidermal cells vary
significantly between species: length varies
from 105.06 ± 16.19 μm (S. grandis, adaxial
surface) to 53.99 ± 7.52 μm (S. intermedia,
abaxial surface); width – from 41.53 ± 3.31 μm
(S. aethiopica) to 13.81 ± 1.13 μm (S. suffruticosa);
height – from 38.99 ± 4.65 μm (S. grandis) to
20.44 ± 3.10 μm (S. gracilis) (Table 1).
The data obtained on the density of
the ordinary epidermal cells demonstrate
significant interspecific variability among
the studied Sansevieria species. In general,
most species exhibit higher cell density on
the abaxial surface, which is probably due
to the functional specialization of the lower
epidermis, particularly its more intensive
participation in transpiration and protection
of the leaf from overheating. The density of
ordinary epidermal cells per unit area (1 mm2)
in the studied species varies more than twofold
(Table 1).
The highest epidermal cell density values
were observed in S. gracilis (996 cells/mm2
on the adaxial surface and 1145 cells/mm2
on the abaxial surface) and S. suffruticosa
(1017 cells/mm² on the abaxial surface). Such
high values may indicate adaptation of these
Plant Introduction • 109 23
Micromorphological traits of the leaf surface in Sansevieria species
A B C
D E F
G H I
J K L
Figure 1. SEM micrographs of Sansevieria leaf surfaces: S. aethiopica (A – adaxial surface, B – abaxial surface);
S. canaliculata (C – abaxial surface); S. cylindrica (D – abaxial surface); S. gracilis (E – adaxial surface, F – abaxial
surface); S. grandis (G – adaxial surface, H – abaxial surface); S. intermedia (I – adaxial surface, J – abaxial
surface); S. kirkii (K – adaxial surface, L – abaxial surface).
species to conditions of increased insolation
and aridity, consistent with their natural
ecology.
In contrast, S. roxburghiana and S. trifasciata
show moderate density values, which may be
related to a wider ecological amplitude and
less dependence on xeromorphic conditions
(Table 1).
The lowest stomatal density values were
observed in S. aethiopica and S. grandis,
which also correlate with the relatively
larger size of their epidermal cells: the larger
the cell area, the fewer stomata per unit
area.
In most species, the abaxial surface
contains more epidermal cells than the
24 Plant Introduction • 109
Maryniuk et al.
adaxial surface. This is a typical feature of
many succulent plants and species with stiff
leaves, in which the lower epidermis plays a
more significant mechanical and regulatory
role: it provides better transpiration
control, greater leaf mechanical strength,
and better thermoregulation. This directly
applies to the studied Sansevieria species,
which are succulent xerophytes. Among
the species we studied, only two (S. grandis
and S. trifasciata) had a higher density of
ordinary epidermal cells on the adaxial
surface. Thus, the analysis clearly illustrates
differences among Sansevieria species in
epidermal cell density and suggests the
existence of species-specific adaptive
strategies related to their anatomical and
ecological peculiarities.
Cuticle and wax deposits
The studied Sansevieria species are
characterized by a well-developed cuticular
cover and the presence of wax deposits
that perform a protective and water-saving
function. The distinctiveness of the cuticular
cover and the details of its external structure
vary both between species and between the
surfaces of leaves of the same species. The
abaxial surface of the leaf blade usually has
a denser cuticular cover compared to the
adaxial surface. This is particularly evident
in S. canaliculata, S. cylindrica, S. gracilis,
S. intermedia, S. kirkii, S. roxburghiana, and
S. suffruticosa (Fig. 1 B, C, F, J, L; Fig. 2 F, G).
Differences are also observed in the
density of the cuticular covering at the
boundary of the ordinary epidermal cells.
Areas of the surface located above the
A B C
D E F
G H I
Figure 2. SEM micrographs of Sansevieria leaf surface: S. metallica (A – adaxial surface, B – abaxial surface);
S. parva (C – adaxial surface, D – abaxial surface); S. roxburghiana (E – adaxial surface, F – abaxial surface);
S. suffruticosa (G – abaxial surface); S. trifasciata (H – adaxial surface, I – abaxial surface).
Plant Introduction • 109 25
Micromorphological traits of the leaf surface in Sansevieria species
Species Cuticle thickness
(µm)
Ordinary epidermal cells sizes (µm) Ordinary
epidermal cells
density (pcs./
mm²)
Adaxial
surface
Abaxial
surface
Adaxial surface Abaxial surface Adaxial
surface
Abaxial
surface
Length
(µm)
Width
(µm)
Height
(µm)
Length
(µm)
Width
(µm)
Height
(µm)
S. aethiopica 5.13 ±
0.65
6.19 ±
0.54
88.89 ±
13.41
41.53 ±
3.31
33.63 ±
3.29
85.31 ±
14.66
33.47 ±
3.77
35.05 ±
3.83
393 ±
18.38
478 ±
82.02
S. cylindrica — 7.89 ±
0.95
— — — 63.93 ±
11.12
18.42 ±
2.02
31.26 ±
4.11
— 888 ±
90.87
S. canaliculata — 7.63 ±
0.84
— — — 60.72 ±
11.05
19.38 ±
2.70
33.52 ±
4.28
— 743 ±
45.40
S. grandis 5.24 ±
0.45
5.26 ±
0.56
105.06
± 16.19
30.34 ±
3.50
38.99 ±
4.65
102.1 ±
21.20
32.91 ±
7.7
31.07 ±
4.34
405 ±
22.76
373 ±
9.98
S. parva 3.39 ±
0.50
3.82 ±
0.69
92.90 ±
20.63
28.90 ±
3.89
32.63 ±
4.06
96.30 ±
14.07
25.30 ±
3.04
22.82 ±
3.48
441 ±
16.26
468 ±
8.49
S. kirkii 7.93 ±
0.69
8.51 ±
0.81
90.93 ±
12.80
22.66 ±
2.31
30.75 ±
3.28
85.85 ±
12.68
21.92 ±
2.72
25.69 ±
2.57
483 ±
25.4
595 ±
30.26
S. roxburghiana 6.31 ±
0.35
8.22 ±
0.63
67.37 ±
9.47
26.39 ±
2.73
33.72 ±
3.08
61.99 ±
11.19
18.47 ±
2.06
30.73 ±
2.94
650 ±
23.15
913 ±
32.47
S. gracilis 7.52 ±
1.31
7.51 ±
0.75
65.90 ±
8.11
19.19 ±
2.18
20.44 ±
3.10
59.38 ±
7.95
17.51 ±
2.25
24.15 ±
3.32
996 ±
124.86
1145 ±
48.81
S. suffruticosa — 11.43 ±
1.71
— — — 57.35 ±
8.84
16.26 ±
2.15
37.40 ±
5.76
— 1017 ±
63.80
S. trifasciata 4.71 ±
0.56
5.87 ±
0.56
74.21 ±
7.79
24.14 ±
2.13
24.83 ±
2.76
71.12 ±
9.28
24.11 ±
2.62
24.22 ±
2.37
706 ±
41.42
648 ±
33.58
S. intermedia 9.23 ±
0.95
9.92 ±
1.20
61.55 ±
11.25
22.01 ±
2.25
36.84 ±
3.41
53.99 ±
7.52
19.35 ±
2.08
37.21 ±
3.35
692 ±
28.15
816 ±
53.29
S. metallica 4.66 ±
0.72
5.60 ±
0.54
99.82 ±
16.73
22.69 ±
2.80
23.34 ±
2.41
101.9 ±
23.07
20.29 ±
3.40
20.20 ±
1.61
528 ±
53.82
671 ±
63.60
Table 1. Morphometric characteristics of the leaf epidermis in the Sansevieria genus.
anticlinal walls are less covered with
wax than the tangential walls of the
cells. In S. cylindrica, S. gracilis, S. kirkii,
S. roxburghiana, and S. suffruticosa, there
are cuticular ridges oriented perpendicular
to the longitudinal rows of epidermal cells
(Fig. 1 C, E, F, K, L; Fig. 2 F, G). In S. gracilis,
the cuticular grooves on the adaxial surface
are slightly less pronounced (Fig. 1 E), while
in S. roxburghiana, cuticular ridges on the
adaxial surface are absent (Fig. 2 E).
The thickness of the cuticle also varies
between species and leaf surfaces. On the
adaxial side, it ranged from 3.39 ± 0.50 μm
(S. parva) to 9.23 ± 0.95 μm (S. intermedia),
and on the abaxial side from 3.82 ± 0.69 μm
(S. parva) to 11.43 ± 1.71 μm (S. suffruticosa)
(Table 1).
Wax deposits on both leaf sides are
represented by a crust (Fig. 3 A–C), a
continuous layer of granular (lumpy) wax
(Fig. 3 D), or a continuous layer (Fig. 3 E).
Crystalloid wax occurs in the form of
irregular crystals (Fig. 3 F–H), occasionally
assembled into rosettes (Fig. 3 A) or in the
form of smooth plates of various shapes and
sizes (Fig. 3 I). Wax crystals can be oriented
perpendicular or at various angles to the
surface of the leaf blade. The crust is mainly
localized on the cells’ surface near stomata,
while wax plates are located around
stomata. The depressions formed by the
immersion of the stomata are also covered
with wax deposits, which is an important
component of the xeromorphic structure
(Fig. 3).
26 Plant Introduction • 109
Maryniuk et al.
Species Stomata sizes (µm) Stomata density
(pcs./mm²)
Stomatal index (%)
Adaxial surface Abaxial surface Adaxial
surface
Abaxial
surface
Adaxial
surface
Abaxial
surface
Length
(µm)
Width
(µm)
Length
(µm)
Width
(µm)
S. aethiopica 26.65 ±
2.90
14.62 ±
1.63
20.88 ±
1.88
13.14 ±
1.47
19 ± 3.08 22 ± 4.94 4.61 4.40
S. cylindrica — — 25.38 ±
1.71
14.14 ±
1.15
— 25 ± 2.09 — 2.73
S. canaliculata — — 33.27 ±
2.23
15.17 ±
1.14
— 9 ± 1.62 — 1.19
S. grandis 29.59 ±
2.49
14.49 ±
1.29
29.54 ±
3.76
16.34 ±
3.34
12 ± 2.41 12 ± 2.39 2.87 3.11
S. parva 34.76 ±
2.31
15.54 ±
2.04
38.52 ±
2.29
13.19 ±
1.74
14 ± 0.82 15 ± 0.96 3.07 3.10
S. kirkii 36.08 ±
2.66
15.70 ±
1.31
31.72 ±
1.88
16.19 ±
1.48
11 ± 1.89 12 ± 1.87 2.22 1.97
S. roxburghiana 22.83 ±
1.88
12.44 ±
0.97
22.59 ±
1.56
12.96 ±
1.00
16 ± 2.08 18 ± 2.88 2.32 1.93
S. gracilis 26.60 ±
1.88
12.83 ±
1.05
25.12 ±
2.09
11.99 ±
1.05
13 ± 2.68 13 ± 2.63 1.28 1.12
S. suffruticosa — — 28.34 ±
2.04
14.74 ±
1.67
— 27 ± 4.02 — 2.58
S. trifasciata 28.28 ±
1.83
19.12 ±
1.97
33.18 ±
2.06
15.92 ±
1.41
12 ± 1.8 12 ± 1.77 1.68 1.67
S. intermedia 31.31 ±
2.81
18.35 ±
1.34
37.55 ±
2.14
18.47 ±
1.50
14 ± 1.74 17 ± 2.45 1.98 2.04
S. metallica 35.71 ±
2.47
14.57 ±
1.17
36.99 ±
2.41
15.70 ±
1.40
14 ± 2.06 14 ± 1.41 2.59 2.05
Table 2. Morphometric characteristics of the leaf stomata in the Sansevieria genus.
Stomatal apparatus
In most of the studied Sansevieria
species, the leaves are amphistomatic,
while hypostomatic leaves are observed
in S. cylindrica, S. canaliculata, and
S. suffruticosa. The stomatal apparatus is
of the anomocytic type: the guard cells of
the stomata are surrounded by subsidiary
cells that do not differ from the rest of the
epidermal cells and have a uniform structure.
The aperture is oriented along the long axis
of the ordinary epidermal cells. Stomata are
unevenly distributed on the surface of the
leaf blade. They occur solitary, in groups of
two or three, or can be organized in rows.
Exclusively solitar stomata are found in
S. parva, while in most other species, both
solitar stomata and groups of two or three
stomata are observed. The most significant
number of stomatal groups is recorded in
S. kirkii on both surfaces of the leaf.
In S. cylindrica, the stomata are located
on the surface of the leaf blade between the
cuticular ridges – stripes, forming more or less
regular groups and rows.
Regarding the location of stomata relative
to the level of the ordinary epidermal cells, the
most deeply immersed stomata are observed
in S. aethiopica, S. cylindrica, S. suffruticosa,
S. grandis, and S. roxburghiana. Stomata of
medium immersion characterize S. gracilis,
S. kirkii, S. metallica, S. intermedia, and
S. canaliculata. In contrast, in S. parva and
S. trifasciata, the stomata are located almost at
the same level as the ordinary epidermal cells
(Fig. 4).
The guard cells of the stomata in most
species, except for S. parva, are surrounded
Plant Introduction • 109 27
Micromorphological traits of the leaf surface in Sansevieria species
A B C
D E F
G H I
Figure 3. SEM micrographs of different types of epicuticular wax on the leaf surface of Sansevieria species:
S. aethiopica (A – adaxial surface); S. metallica (B – adaxial surface); S. cylindrica (C – abaxial surface);
S. suffruticosa (D – abaxial surface); S. parva (E – adaxial surface); S. grandis (F – adaxial surface); S. aethiopica
(G – adaxial surface); S. intermedia (H – abaxial surface); S. trifasciata (I – adaxial surface).
by a distinct cuticular ridge of rounded or
rectangular shape (Fig. 4), a characteristic
mainly found in xerophytic plants. The cuticle
of these ridges is significantly thickened
and raised above the general surface of the
epidermis. In S. parva, the cuticular ring is
absent. All species also have large sub-stomatal
cavities (Fig. 4).
The data obtained indicate that the density
of stomata on the adaxial and abaxial sides of
the leaf in all studied species is low, ranging
from 9 to 27 per 1 mm2. The highest number of
stomata per unit area is found in S. cylindrica,
S. suffruticosa, and S. aethiopica, which have
cylindrical (functionally abaxial) or vertically
oriented leaves. The highest density of
stomata compared to other species was found
in S. suffruticosa (27 ± 4.02 per 1 mm2), and the
lowest in S. canaliculata (9 ± 1.62 per 1 mm2)
(Table 2).
In the studied species with amphistomatic
leaf blades, there is no clear predominance of
stomata on the adaxial or abaxial surface. Both
sides of the leaf have almost the same stomatal
density. This is probably due to the specific
position of the leaf blade, which is vertical in
most species. As a result, approximately the
same amount of solar radiation falls on the
upper and lower surfaces of the leaf, which
leads to a relatively uniform distribution
of stomata on both sides to optimize gas
exchange and minimize water loss.
The length of the stomata varies from
38.52 ± 2.29 μm (S. parva, adaxial surface)
to 20.88 ± 1.88 μm (S. aethiopica, abaxial
surface). The width of the stomata varies from
28 Plant Introduction • 109
Maryniuk et al.
A1 A2 B1 B2
C1 C2 D1 D2
E1 E2 F1 F2
G1 G2 H1 H2
I1 I2 J1 J2
K1 K2 L1 L2
Figure 4. Microstructure of Sansevieria stomata on a leaf cross-sections (1) and their SEM micrographs (2):
S. aethiopica (A); S. canaliculata (B); S. cylindrica (C); S. gracilis (D); S. grandis (E); S. intermedia (F); S. kirkii (G);
S. metallica (H); S. parva (I); S. roxburghiana (J); S. suffruticosa (K); S. trifasciata (L).
Plant Introduction • 109 29
Micromorphological traits of the leaf surface in Sansevieria species
19.12 ± 1.97 μm (S. trifasciata, adaxial surface)
to 11.99 ± 1.05 μm (S. gracilis, abaxial surface)
(Table 2).
There were also cases in which stomata
formed clusters, contacting each other, mainly
laterally (Fig. 4 A1, B1, K1).
Our research has revealed that some of
the stomata on the leaf surface of the studied
Sansevieria species remain underdeveloped
(Fig. 1 A, B, G; Fig. 2 E, F, H, I). We observed
underdevelopment of stomata in all species
studied, except S. suffruticosa, S. cylindrica,
and S. canaliculata. In S. roxburghiana and
S. trifasciata, more than half of the stomata
remain underdeveloped; however, no apparent
connection with the location on the surface
(adaxial or abaxial) was observed. These results
are entirely consistent with the data from the
experimental work of Kagan & Sachs (1991),
which states that in S. trifasciata up to half
of the potential stomata remain ‘immature’.
Kagan & Sachs (1991) remains the most cited
on the phenomenon of ‘immature’ stomata
in Sansevieria and examines the mechanism
of development and spatial distribution of
mature/immature stomata.
Stomatal index
The stomatal index varies from 4.4
(S. aethiopica) to 1.12 (S. gracilis). Low values
correspond to pronounced xeromorphic traits
and a higher epidermal cell density.
Discussion
The data obtained indicate that representatives
of the Sansevieria genus exhibit a wide
range of anatomical and micromorphological
adaptations to minimize water loss and
optimize gas exchange in arid environments.
The most informative features are obviously
the thickness of the cuticle, the type of wax
deposits, the density of stomata and the
degree of their immersion, the stomatal
index, and the ratio of the sizes and density of
epidermal cells.
Certain data on the Sansevieria genus we
found in this study contradict the reports
of other authors. In particular, Shewale
et al. (2022) and Jasna et al. (2025) reported
the presence of unicellular trichomes in
S. cylindrica, but trichomes were not observed
during our research.
Considering that most Sansevieria species
grow in environments with periodic droughts
but without intense ultraviolet radiation,
i.e., often under the cover of sparse forest
(Takawira & Nordal, 2002; Burkart et al., 2025),
the absence of trichomes is a justifiable and
evolutionarily argued trait.
Unlike other succulent plants, in which
trichomes act as structures that reduce
evaporation or scatter excess light,
Sansevieria species implement a different
xeromorphic strategy: thickening of the
cuticle and development of epicuticular
wax (Klimko et al., 2018; Maděra et al., 2020),
pronounced depression of stomata (Klimko
et al., 2018), and the presence of thick-walled
epidermal cells (Maděra et al., 2020). Such a
set of characteristics compensates for the
functional significance of trichomes, since
other morphological mechanisms ensure
water retention capacity.
Data on the density and size of ordinary
epidermal cells in the studied Sansevieria
species reflect a compromise between
epidermal cell size and density that is universal
for plants: as cell size decreases, their number
per unit area increases, consistent with the
results (John et al., 2017). This pattern is due to
structural limitations of the tissue and is well
described for leaf epidermis. The increased
cell density on the abaxial surface, observed
in most of the species studied, corresponds to
the general anatomical trends characteristic
of xerophytic plants (Fahn, 2002; Ogburn &
Edwards, 2010), and can be considered an
adaptive feature associated with reduced
transpiration losses and enhanced mechanical
stability of the leaf.
The presence of cuticular ridges and
variability in cuticle thickness reflects
interspecific differences in adaptation
strategies. Higher values of cuticle density and
thickness on the abaxial surface may provide
additional protection against water loss and
mechanical damage. In contrast, a less dense
cuticular covering on the adaxial surface
may promote optimal gas exchange through
stomata.
The species S. aethiopica, S. cylindrica,
and S. suffruticosa are characterized by a
complex of pronounced xeromorphic features:
thickened cuticle, deeply recessed stomata,
high density of wax deposits, and low stomatal
index. This is consistent with their ecology, as
30 Plant Introduction • 109
Maryniuk et al.
these species grow in regions with minimal
rainfall on dry rocky slopes and in seasonally
dry tropical biomes with prolonged drought
(POWO, 2025). In contrast, S. canaliculata
exhibits a different adaptive model: low
stomatal density, but larger stomata and
shallower stomatal depth. This combination
may be related to the optimization of the ratio
of transpiration to nighttime CO2 uptake in
plants with CAM metabolism (Martin et al.,
2019).
In species with vertically oriented or
cylindrical leaf blades (i.e., S. intermedia,
S. metallica, S. grandis), there is no significant
predominance of stomata on any of the
surfaces. However, most angiosperms are
characterized by a predominance of stomata
on the abaxial surface of amphistomatic
leaves (Fahn, 2002). Most probably, in the
above-mentioned Sansevieria species, the
violation of this pattern is explained by the
uniform illumination of both sides of the leaf.
These species also tend to increase epidermal
cell size while simultaneously reducing cell
density, suggesting species-specific strategies
to minimize evaporation.
Amphistomatousness is characteristic
of Sansevieria species with bifacial leaves
(i.e., S. aethiopica, S. trifasciata) and, in some
species, can be interpreted as an adaptive trait
to intense illumination. It is combined with
changes in the number and size of ordinary
epidermal cells, which ensure a more uniform
distribution of light flux and stable gas
exchange.
Leaf stomata are not only an essential
regulatory element that ensures a balance
between photosynthesis and transpiration,
but also a key structure that influences
atmospheric carbon and water cycles
(Hetherington & Woodward, 2003). Stomatal
density and size are important traits that show
significant variability among plant species in
response to environmental factors such as
temperature, light intensity, water content,
soil nutrient levels, and atmospheric CO2
concentration (Dow et al., 2014; Durand et al.,
2020).
The anomocytic stomata we found in all
studied Sansevieria species, according to Rudal
et al. (2017), probably represent the original
(plesiomorphic) state in monocots, although
many reversions have occurred during their
evolution. However, Babu & Srinivasa Prabhu
(2024), in their comprehensive investigation
of S. trifasciata, refer to the tetracytic type of
stomata in this species.
The presence of ‘immature’ stomata is
interpreted as a developmental reserve
that allows individual cells to be potentially
activated or completed depending on the
environment, especially in young leaves (Han
et al., 2021). Underdeveloped stomata do not
seem to be a random defect, but the result of
specific regulation of epidermal development.
In many plants, including S. trifasciata, some
stomata fail to develop at an early stage. This
has several biological reasons and can be
considered an adaptive phenomenon.
In this context, the presence of
underdeveloped stomata may indicate the
genus’s adaptive capabilities. For Sansevieria,
which naturally exists in conditions of periodic
drought, high-intensity lighting, and limited
water availability, such a system of stomatal
development offers several advantages:
water conservation, tissue stability during
fluctuations in moisture, maintenance of CAM
photosynthesis, and the ability to respond
quickly and flexibly to changes in growing
conditions. In other words, underdeveloped
stomata are not only a result of development
but also a structural feature that contributes
to Sansevieria high ecological resilience and
its ability to survive in arid environments or
conditions of introduction (Kagan & Sachs,
1991).
The stomatal index proved to be an
informative diagnostic parameter that enables
comparison of adaptive gas exchange models
across closely related taxa.
Plants of S. gracilis and S. roxburghiana,
which grow in less extreme conditions, are
characterized by less pronounced xeromorphic
features – an increase in the size of epidermal
cells, which corresponds to their ecological
niche. Mesomorphic traits are also found in
S. parva – its stomata are located almost at the
same level as the epidermal cells, and there is
no cuticular roller around the stomata. This
indicates adaptation to conditions with a less
pronounced moisture deficit.
The functional relationship between leaf
surface microstructures and the maintenance
of water balance was also demonstrated in
a study by Jura-Morawiec & Marcinkiewicz
(2020) on Dracaena draco (L.) L. The authors
showed that micromorphological features
Plant Introduction • 109 31
Micromorphological traits of the leaf surface in Sansevieria species
of the leaf surface affect wettability, water
absorption, and the ability to store water
temporarily. These properties are directly
related to the micromorphology of the cuticle
and wax layer and reflect adaptations to arid
conditions.
Conclusions
Hence, analysis of the micromorphological
features of the epidermis of the studied
Sansevieria species revealed a complex of
xeromorphic traits consistent with their
ecological survival strategies in arid conditions.
At the level of leaf micromorphology,
amphistomatousness, spatial organization
of epidermal cells, the cuticular cover
with epicuticular wax deposits of various
configurations, varying degrees of stomatal
immersion, and variation in their density,
as well as the presence of underdeveloped
stomata.
The interspecific differences identified
indicate distinct ways of optimizing water
balance, which are shaped by similar but not
identical growing conditions. In general, this
study can be considered a model, and its
results can be extrapolated to other members
of Asparagaceae and Asparagales.
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Plant Introduction • 109 33
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Мікроморфологічні особливості поверхні листка видів роду Sansevieriа Thunb.
s.str. (Asparagaceae)
Мирослава Маринюк *, Людмила Буюн, Олександр Гиренко, Іван Гурненко
Національний ботанічний сад імені М.М. Гришка НАН України, вул. Садово-Ботанічна, 1, Київ, 01103,
Україна; * maryniukmyroslava@gmail.com
У дослідженні за допомогою світлової мікроскопії та сканувальної електронної мікроскопії вивчено
мікроморфологію поверхні листків 12 видів роду Sansevieria. Основну увагу зосереджено на виявленні
мікроморфологічних ознак, пов’язаних зі стійкістю рослин до стресових умов, зокрема форми клітин
епідерми, товщини кутикули, розподілу та щільності продихів, а також особливостей кутикулярного
покриву з епікутикулярними восковими відкладеннями. У більшості досліджених видів Sansevieria
листки є амфістоматичними, тоді як гіпостоматичні листки виявлено у S. cylindrica, S. canaliculata та
S. suffruticosa. У всіх досліджених таксонів епідерма представлена одним шаром клітин і позбавлена
трихом.
Рослини досліджених видів Sansevieria характеризуються добре розвиненим кутикулярним шаром і
наявністю воскових відкладень, що виконують захисну та водозберігаючу функції. Ступінь розвитку
кутикули та її структура варіює як між видами, так і між поверхнями листка в межах одного виду.
Абаксіальна поверхня листкової пластинки зазвичай має більш розвинений кутикулярний покрив
порівняно з адаксіальною, що особливо чітко проявляється у S. cylindrica, S. canaliculata, S. kirkii,
S. roxburghiana, S. gracilis, S. suffruticosa та S. intermedia. Усі досліджені види мають аномоцитні продихи.
Щільність продихів на абаксіальній поверхні листка коливалася в межах 9–27 мм2 залежно від виду.
Виявлені міжвидові відмінності відображають різні стратегії оптимізації водного балансу за аридних
умов.
https://doi.org/10.11646/phytotaxa.376.6.2
https://doi.org/10.11646/phytotaxa.376.6.2
https://doi.org/10.21123/bsj.2017.14.1.0189
https://doi.org/10.1111/boj.12385
https://doi.org/10.3390/molecules25112608
https://agrobiodiversity.uniag.sk/scientificpapers/article/view/119
https://agrobiodiversity.uniag.sk/scientificpapers/article/view/119
https://doi.org/10.1016/j.ympev.2022.107404
https://doi.org/10.1016/j.ympev.2022.107404
https://doi.org/10.3897/phytokeys.185.72259
https://doi.org/10.5281/zenodo.2330922
https://doi.org/10.5281/zenodo.2330922
34 Plant Introduction • 109
Maryniuk et al.
На рівні мікроморфології листка амфістоматичність, просторова організація клітин епідерми,
наявність кутикулярного покриву з епікутикулярними восковими відкладеннями різної конфігурації,
різний ступінь занурення та щільності продихів, а також наявність недорозвинених продихів
можуть розглядатися як маркери стійкості рослин цього роду до стресових факторів. Виявлені
мікроморфологічні маркери поглиблюють уявлення про адаптивні ксероморфні ознаки Sansevieria
та мають потенційне прикладне значення для біотехнологічних розробок і фіторемедіації, зокрема
для розбудови зеленої інфраструктури.
Ключові слова: Sansevieria, світлова мікроскопія, сканувальна електронна мікроскопія, кутикула, епідерма, продиховий
апарат, мікроморфологія, фіторемедіація
|
| id | oai:ojs2.plantintroduction.org:article-1683 |
| institution | Plant Introduction |
| keywords_txt_mv | keywords |
| language | English |
| last_indexed | 2026-02-08T08:11:58Z |
| publishDate | 2026 |
| publisher | M.M. Gryshko National Botanical Garden of the NAS of Ukraine |
| record_format | ojs |
| resource_txt_mv | wwwplantintroductionorg/fe/69b655df39477f7536d4b3912ced07fe.pdf |
| spelling | oai:ojs2.plantintroduction.org:article-16832026-01-19T02:05:33Z Micromorphological traits of the leaf surface in species of the genus Sansevieria Thunb. s.str. (Asparagaceae) Мікроморфологічні особливості поверхні листка видів роду Sansevieriа Thunb. s.str. (Asparagaceae) Maryniuk, Myroslava Buyun, Lyudmyla Gyrenko, Oleksandr Gurnenko, Ivan This study investigated the leaf surface micromorphology of 12 species of the genus Sansevieria using light microscopy and scanning electron microscopy. The research focused on identifying micromorphological traits associated with plant stress tolerance, including epidermal cell shape, cuticle thickness, stomatal distribution and density, and cuticular characteristics with epicuticular wax deposits. In most of the studied Sansevieria species, the leaves are amphistomatic, whereas hypostomatic leaves are observed in S. cylindrica, S. canaliculata, and S. suffruticosa. In all studied taxa, the epidermis consists of a single layer of cells and lacks trichomes.The examined Sansevieria species are characterized by a well-developed cuticular layer and the presence of wax deposits that perform protective and water-conserving functions. The thickness of the cuticle and its ornamentation vary both among species and between leaf surfaces within the same species. The abaxial leaf surface generally exhibits a more developed cuticle than the adaxial surface, a feature particularly pronounced in S. cylindrica, S. canaliculata, S. kirkii, S. roxburghiana, S. gracilis, S. suffruticosa, and S. intermedia. All investigated species possess anomocytic stomata. Stomatal density on the abaxial leaf surface ranged from 9 to 27 mm2 among the studied species. These interspecific variations reflect distinct strategies for optimizing water balance under arid conditions.At the level of leaf micromorphology, amphistomaty, the spatial organization of epidermal cells, the presence of a cuticular layer with epicuticular wax deposits of various configurations, differences in stomatal sunkenness and density, and the occurrence of underdeveloped stomata can be considered markers of stress tolerance in this genus. The identified micromorphological markers provide insight into the adaptive xeromorphic traits of Sansevieria and have potential applications in applied research, including biotechnological projects and phytoremediation, including green infrastructure development. У дослідженні за допомогою світлової мікроскопії та сканувальної електронної мікроскопії вивчено мікроморфологію поверхні листків 12 видів роду Sansevieria. Основну увагу зосереджено на виявленні мікроморфологічних ознак, пов’язаних зі стійкістю рослин до стресових умов, зокрема форми клітин епідерми, товщини кутикули, розподілу та щільності продихів, а також особливостей кутикулярного покриву з епікутикулярними восковими відкладеннями. У більшості досліджених видів Sansevieria листки є амфістоматичними, тоді як гіпостоматичні листки виявлено у S. cylindrica, S. canaliculata та S. suffruticosa. У всіх досліджених таксонів епідерма представлена одним шаром клітин і позбавлена трихом.Рослини досліджених видів Sansevieria характеризуються добре розвиненим кутикулярним шаром і наявністю воскових відкладень, що виконують захисну та водозберігаючу функції. Ступінь розвитку кутикули та її структура варіює як між видами, так і між поверхнями листка в межах одного виду. Абаксіальна поверхня листкової пластинки зазвичай має більш розвинений кутикулярний покрив порівняно з адаксіальною, що особливо чітко проявляється у S. cylindrica, S. canaliculata, S. kirkii, S. roxburghiana, S. gracilis, S. suffruticosa та S. intermedia. Усі досліджені види мають аномоцитні продихи. Щільність продихів на абаксіальній поверхні листка коливалася в межах 9–27 мм2 залежно від виду. Виявлені міжвидові відмінності відображають різні стратегії оптимізації водного балансу за аридних умов. На рівні мікроморфології листка амфістоматичність, просторова організація клітин епідерми, наявність кутикулярного покриву з епікутикулярними восковими відкладеннями різної конфігурації, різний ступінь занурення та щільності продихів, а також наявність недорозвинених продихів можуть розглядатися як маркери стійкості рослин цього роду до стресових факторів. Виявлені мікроморфологічні маркери поглиблюють уявлення про адаптивні ксероморфні ознаки Sansevieria та мають потенційне прикладне значення для біотехнологічних розробок і фіторемедіації, зокрема для розбудови зеленої інфраструктури. M.M. Gryshko National Botanical Garden of the NAS of Ukraine 2026-01-18 Article Article application/pdf https://www.plantintroduction.org/index.php/pi/article/view/1683 10.46341/PI2025024 Plant Introduction; No 109 (2026): Early view; 20-34 Інтродукція Рослин; № 109 (2026): Early view; 20-34 2663-290X 1605-6574 en https://www.plantintroduction.org/index.php/pi/article/view/1683/1581 Copyright (c) 2026 Myroslava Maryniuk, Lyudmyla Buyun, Oleksandr Gyrenko, Ivan Gurnenko http://creativecommons.org/licenses/by/4.0 |
| spellingShingle | Maryniuk, Myroslava Buyun, Lyudmyla Gyrenko, Oleksandr Gurnenko, Ivan Мікроморфологічні особливості поверхні листка видів роду Sansevieriа Thunb. s.str. (Asparagaceae) |
| title | Мікроморфологічні особливості поверхні листка видів роду Sansevieriа Thunb. s.str. (Asparagaceae) |
| title_alt | Micromorphological traits of the leaf surface in species of the genus Sansevieria Thunb. s.str. (Asparagaceae) |
| title_full | Мікроморфологічні особливості поверхні листка видів роду Sansevieriа Thunb. s.str. (Asparagaceae) |
| title_fullStr | Мікроморфологічні особливості поверхні листка видів роду Sansevieriа Thunb. s.str. (Asparagaceae) |
| title_full_unstemmed | Мікроморфологічні особливості поверхні листка видів роду Sansevieriа Thunb. s.str. (Asparagaceae) |
| title_short | Мікроморфологічні особливості поверхні листка видів роду Sansevieriа Thunb. s.str. (Asparagaceae) |
| title_sort | мікроморфологічні особливості поверхні листка видів роду sansevieriа thunb. s.str. (asparagaceae) |
| url | https://www.plantintroduction.org/index.php/pi/article/view/1683 |
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