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RESEARCH ARTICLE
Taxonomic significance of seed macro-micromorphology of
Turkish Alcea L. (Malvaceae) through light microscopy and
scanning electron microscopy
Funda Özbek | Mehmet Erkan Uzunhisarcıklı
Department of Biology, Faculty of Science, Gazi University, Ankara, Turkey
Correspondence
Funda Özbek, Department of Biology, Faculty
of Science, Gazi University, 06500,
Teknikokullar/Ankara, Turkey.
Email: fundaozbek@gazi.edu.tr
Funding information
Scientific and Technological Research Council
of Turkey for the collection of plant materials,
Grant/Award Number: TUBITAK-TBAG 2282
Review Editor: Paul Verkade
Abstract
Seed morphological properties of 19 taxa belonging to the genus Alcea L. (Malvaceae)
distributed in Turkey were investigated using a light microscope and scanning elec-
tron microscope to identify their characters and to evaluate their diagnostic value.
The seeds are reniform with a rounded apex and base, reniform in shape, and light to
dark brown, grayish-brown, or blackish-brown in color. The seed length ranges from
2.22 to 6.5 mm and seed width from 1.72 to 6.5 mm. The indumentum at the ventral
and dorsal regions of the seed differs in density. Three types of seed coat ornamenta-
tions were observed: reticulate, reticulate-rugulate, and reticulate-ruminate on the
dorsal and lateral faces. Principal component analysis was used to evaluate the impor-
tant seed morphological characteristics among the taxa studied, with four compo-
nents accounting for 90.761% of the total variance. Numerical analysis revealed that
seed size, color, seed surface patterns on dorsal and lateral sides, indumentum at dor-
sal and ventral regions, and periclinal surface sculpture of epidermal cells are particu-
larly the most useful variables for discriminating the Alcea taxa. The findings also
showed a partial relationship among the Alcea taxa clusters, based on seed morphol-
ogy and the systematics of these taxa, based on general macromorphology. Taxo-
nomic key using the seed features is provided to identify the species studied. The
current work will contribute to the knowledge about the family Malvaceae, and
microscopic macro-micromorphological analysis can be used for identification by the
taxonomists for further studies on this family.
Research Highlights
•Seed color, indumentum and surface sculpturing have systematic value for sepa-
rating the taxa.
•Seed morphology of the Alcea taxa was studied via light microscope and scanning
electron microscope.
•Numerical analysis provided the contribution of seed characters to taxa relationships.
Received: 30 January 2023 Revised: 19 May 2023 Accepted: 18 June 2023
DOI: 10.1002/jemt.24385
This is an open access article under the terms of the Creative Commons Attribution-NonCommercial-NoDerivs License, which permits use and distribution in any
medium, provided the original work is properly cited, the use is non-commercial and no modifications or adaptations are made.
© 2023 The Authors. Microscopy Research and Technique published by Wiley Periodicals LLC.
Microsc Res Tech. 2023;1–17. wileyonlinelibrary.com/journal/jemt 1
KEYWORDS
Alcea, identification, numerical analysis, seed, systematics
1|INTRODUCTION
The family Malvaceae Juss. (s.l.) is commonly known as the “Mallow
family”, which contains 4225 species in 224 genera comprised of
herbs, shrubs and small trees. This family is represented by 15 genera
and 55 species in Turkey. Its members are distributed throughout the
world, and mainly occur in warm temperatures and tropical countries,
excluding the cold regions. The genus Alcea L. is taxonomically attrib-
uted to the tribe Malveae, and subfamily Malvoideae of the Malva-
ceae. It comprises about 70 taxa widely distributed throughout
Europe (except for the northern part), North America, North Africa,
Caucasus, Southwest and Central Asia, Southern Russia, and
Afghanistan (Christenhusz & Byng, 2016; Cullen, 1967;
Heywood, 1978; Hutchinson, 1964; Tamde et al., 2016; Uzunhisarcıklı
et al., 2012; Uzunhisarcıklı& Vural, 2012;Yıldırım et al., 2019). The
name of this genus is derived either from alce (remedy) in reference to
its therapeutic value, or from alke (strength), because of its vigorous
growth (Fryxell, 1997).
Alcea is represented by 18 species in the Flora of Turkey
(Cullen, 1967). Of these species, four are cited as endemic species.
After the revision of this genus, Alcea lavateriflora (DC.) Boiss. was
considered a doubtful record by Uzunhisarcıklıand Vural (2012), and
they reported A. rosea L. as a new cultivated record. However, these
researchers stated that Alcea apterocarpa (Fenzl) Boiss., Alcea calvertii
(Boiss.) Boiss. and Alcea fasciculiflora Zohary have been reported in
neighboring countries, and so these species are not regarded as
endemic to Turkey. At present, this genus consists of 19 taxa (17 spe-
cies, one subspecies and one cultural species), and only Alcea pisidica
Hub.-Mor. is endemic to Turkey (Uzunhisarcıklı& Vural, 2012).
In the taxonomic history of the genus Alcea, Linnaeus (1753) ear-
lier evaluated both Alcea and Althaea L. as separate genera. In the sub-
sequent studies, Willdenow (1800), De Candolle (1824), and Bentham
and Hooker (1862) fused the species of two genera into one genus,
Althaea. Later, Cullen (1967) adopted the idea accepted by many tax-
onomists such as Alefeld (1862), Boissier (1867), and Iljin (1949)to
treat both Althaea and Alcea as two distinct genera based on their car-
pel and anther features.
The significance of seed morphological characteristics in plant
systematics for the family Malvaceae has been emphasized in some
studies (Ahmed & Qaiser, 1989; Ather et al., 2009; Kirkbride
et al., 2006; Masullo et al., 2020; Özbek & Uzunhisarcıklı,2020a;
2020b; Sivarajan & Pradeep, 1996). Paul and Nayar (1987) examined
the seeds of 54 taxa under 17 genera using SEM, and said that the
variation in the seed coat surface differs from species to species, even
in infraspecific ranks. El Naggar (2001) investigated the seed morpho-
logical properties of 14 Egyptian species, and these taxa were divided
into two groups based on anticlinal cell boundaries and periclinal cell
wall sculpture. Esteves (2004) stated that the seeds have taxonomical
significance, especially at sectional levels, and the sculpturing of the
seed coat is very important for separating the sections of the subge-
nera Pavonia Cav. Ather et al. (2009) detected that seed macro- and
micromorphological features have a diagnostic value in delimitation of
the taxa at generic and specific ranks. El-Kholy et al. (2011) observed
four types of seed surface ornamentations, and they stated that the
usage of SEM in the examination of the seed coat of 11 cultivars
belonging to two Hibiscus L. species revealed the importance of this
technique as a good taxonomic tool. The seed morphology of the
genus Abelmoschus Medik. was studied, and especially seed shape and
trichome structure were found to be valuable characteristics for sepa-
rating taxa by Patil et al. (2015). Abid et al. (2016) investigated the
seed morphological properties of 75 taxa attributed to Malvaceae
growing in Pakistan using LM and SEM, and highlighted that these
data are taxonomically significant in discrimination of the taxa. Fawzi
(2018) examined five Corchorus L. species for their macro- and micro-
morphological characteristics in seeds, and reported that these prop-
erties are quite diverse and stable so they can be useful for
delimitation of the taxa at specific levels in the species investigated.
Pour et al. (2019) detected that seed shape, surface, indumentum,
strophiole, and seed size are significant for the delimitation of genera
and species in some selected Malvaceae species. Abdel Khalik et al.
(2021) observed four main categories of seed surface: reticulate, fove-
ate, alveolate, and rugose. They stated that seed coat analysis pro-
vided useful information for evaluating the taxonomy of Malvoideae
at both intrageneric and tribal levels.
This research incorporates detailed seed macro- and micromor-
phological characteristics of 19 taxa of the genus Alcea, which are dis-
tributed in Turkey. No studies have been conducted for the seed
morphological features of this genus from Turkey. The objectives of
this study are to provide the seed morphology of this genus via LM
and SEM, and to assess the diagnostic value of these properties for
the taxonomic contribution to the genus.
2|MATERIALS AND METHODS
2.1 |Plant materials
The present study was based on available materials, which were col-
lected from their type location and different areas of Turkey at flow-
ering and fruiting periods between 2003 and 2007 during the revision
of the genus within the scope of the Scientific and Technological
Research Council of Turkey (TUBITAK, Project No: TBAG-2282), and
some plant specimens were acquired from the Prof. Dr. Tuna Ekim
Herbarium (GAZI) at the Gazi University, Faculty of Science. Voucher
numbers, locality information, and the distribution map of the speci-
mens investigated were given in Table 1and Figure 1. The authors of
2ÖZBEK and UZUNHISARCIKLI
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TABLE 1 List of the investigated Alcea L. specimens with location data and voucher number.
Taxa Locality Voucher number
A1 Alcea acaulis (Cav.) Alef. C7 Şanlıurfa: Şanlıurfa-Akçakale, 29 km, 400 m,
13.06.2004, fields
C6 Gaziantep: Nizip-Osmaniye, 300 m, 08.05.2007,
roadsides
M.E. Uzunhisarcıklı1958
M.E. Uzunhisarcıklı2224
A2 A. striata (DC.) Alef. subsp. striata C3 Alanya: Antalya-Isparta, 55 km, 200 m, 06.07.2005,
roadsides
C5 Adana: Saimbeyli-Feke, 27 km to Feke, 800 m,
08.07.2005, roadsides
M.E. Uzunhisarcıklı2039
M.E. Uzunhisarcıklı2068
A3 A. striata (DC.) Alef. subsp.
rufescens (Boiss.) Cullen
B7 Malatya: Malatya-Pötürge, 2 km, 1245 m,
11.07.2005, roadsides
M.E. Uzunhisarcıklı2084
A4 A. remotiflora (Boiss. & Heldr.) Alef. C3 Antalya: Antalya-Isparta, 35 km, 160 m,
06.07.2005, roadsides
C5 Adana: Saimbeyli-Tufanbeyli, 3 km, 1200 m,
08.07.2005, calcareous
M.E. Uzunhisarcıklı2040
M.E. Uzunhisarcıklı2056
A5 A. digitata (Boiss.) Alef. C6 Gaziantep: Gaziantep-Nizip, 10 km, 800 m,
09.07.2005, roadsides
C3 Antalya: Isparta-Antalya, 25 km to Antalya, 125 m,
06.07.2005, roadsides
M.E. Uzunhisarcıklı2074
M.E. Uzunhisarcıklı2042
A6 A. setosa (Boiss.) Alef. C7 Mardin: Nusaybin-Mardin, 48 km, 600 m,
12.06.2004, rocky slopes
C6 Maras¸: Engizek D., around Aksu District, 1000 m,
11.06.1987, fields
M.E. Uzunhisarcıklı1957
H. Duman 3090
A7 A. apterocarpa (Fenzl) Boiss. B8 Erzurum: Pasinler-Hınıs, 29 km to Hınıs, 1875 m,
19.08.2005, roadsides
C2 Denizli: Dazkırı-Denizli, Acıgöl, 850 m, 22.06.2003,
roadsides
M.E. Uzunhisarcıklı2117
M.E. Uzunhisarcıklı1886
A8 A. kurdica (Schltdl.) Alef. C10 Hakkari: Van-Hakkari, 60 km to Hakkari, 1830 m,
17.08.2005, calcareous
B9 Van: Van-Ercis¸, 40 km to Ercis¸, 1800–2000 m,
17.10.1992, steppe, roadsides
M.E. Uzunhisarcıklı2101
H. Özçelik & Y. Altan 4743
A9 A. heldreichii (Boiss.) Boiss. C3 Antalya: Antalya-Isparta, 20 km, 150 m,
06.07.2005, roadsides
B4 Ankara: Polatlı,Sıratas¸lar Tepesi, 840–850 m,
02.09.2004, steppe
M.E. Uzunhisarcıklı2041
M.E. Uzunhisarcıklı2016
A10 A. calvertii (Boiss.) Boiss. A9 Kars: Göle-Şenkaya, 18 km from Şenkaya, 1360 m,
19.08.2005, stony slopes
B8 Erzurum: As¸kale-Tercan, 15–20. km, 1475 m,
20.08.2005, roadsides
M.E. Uzunhisarcıklı2113
M.E. Uzunhisarcıklı2119
A11 A. hohenackeri Boiss. A9 Erzurum: Erzurum-Kars, 101 km, 1541 m,
16.08.2004, roadsides
B7 Diyarbakır: Ergani-Maden, 10 km to Maden, 830 m,
11.07.2005, roadsides
M.E. Uzunhisarcıklı2000
M.E. Uzunhisarcıklı2082
A12 A. pisidica Hub.-Mor.
a
B3 Isparta: Gelendost-E
girdir, 43 km to E
girdir, 936 m,
08.07.2004, fallow field
M.E. Uzunhisarcıklı1987
A13 A. guestii Zohary C7 Şanlıurfa: Virans¸ehir-Ceylanpınar, 34–36 km,
450 m, 12.06.2004, roadsides, steppe
M.E. Uzunhisarcıklı1954
A14 A. biennis Winterl A4 Ankara: Nallıhan-Beypazarı, 28 km, 500–550 m,
15.06.2004, roadsides
C3 Antalya: Manavgat-Akseki, 465 m, 06.07.2005,
roadsides
M.E. Uzunhisarcıklı1973
M.E. Uzunhisarcıklı2046
A15 A. dissecta (Baker f.) Zohary C7 Şanlıurfa: Şanlıurfa-Virans¸ehir, Çobanbo
gazı, 38 km,
480 m, 12.06.2004, roadsides
B7 Elazı
g: Elazı
g-Bingöl, 40 km to Bingöl, 1590 m,
16.08.2005, calcareous
M.E. Uzunhisarcıklı1958
M.E. Uzunhisarcıklı2098
A16 A. excubita Iljin A9 Artvin: Artvin-Ardanuç, 10–15 km, 600 m,
17.08.2004, roadside, stony slopes
B7 Malatya: Malatya-Yazıhan, 13 km to Yazıhan,
720 m, 12.07.2005, roadsides
M.E. Uzunhisarcıklı2013
M.E. Uzunhisarcıklı2086
(Continues)
ÖZBEK and UZUNHISARCIKLI 3
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plant names were written according to Plants of the World Online
(POWO, 2023). The order of the species was adopted from Uzunhi-
sarcıklıand Vural (2012).
2.2 |Light microscopic analysis
Mature seeds were used in the analysis of seed macro- and micromor-
phological characteristics. Seed's shape, minimum and maximum
lengths, minimum and maximum widths, color, indumentum density at
dorsal and ventral regions, and hair type and color were investigated
using a stereoscopic microscope (Leica EZ4D, Wetzlar, Germany). At
least 30 seeds per species were measured for their lengths and widths
for determining the variation among species. Photographs of seeds
were taken with a stereomicroscope digital photomicrograph system
(Olympus SZX7, Tokyo, Japan).
2.3 |Scanning electron microscopic analysis
Micromorphological features of seeds were analyzed through SEM.
First, seeds were cleaned with a mixture of ethanol and xylol (1:1,
v/v). Subsequently, the seeds were transferred onto aluminum stubs
using double-sided adhesive tapes, with their lateral and dorsal sides
TABLE 1 (Continued)
Taxa Locality Voucher number
A17 A. flavovirens (Boiss. & Buhse) Iljin B10 I
gdır: I
gdır-Do
gubeyazıt, 25–30 km, 1400–
1450 m, 30.07.2006, volcanic lands
C10 Hakkari: Hakkari-Van, 40 km, 1500 m,
16.07.2001, stony slopes
M.E. Uzunhisarcıklı2212
Z. Aytaç 8209
A18 A. fasciculiflora Zohary B8 Diyarbakır: Kozluk-Bitlis, 3 km, 680 m, 10.07.2005,
roadsides
M.E. Uzunhisarcıklı2078
A19 A. rosea L. C3 Antalya: Perge, 20 m, 08.07.2004, roadsides
B4 Ankara: AyvalıDistrict, Sahrayıcedit Street, 925 m,
03.09.2021, roadsides
M.E. Uzunhisarcıklı1989a
F. Özbek 1054
a
Endemic taxa.
FIGURE 1 The studied localities of Alcea L. in Turkey.
4ÖZBEK and UZUNHISARCIKLI
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visible, and coated with gold in a sputter coater (Polaron SC 502, Lon-
don, England). They were investigated and photographed with SEM
(JEOL JSM 6060 LV, Tokyo, Japan) at a voltage of 15 kV in the Prof.
Dr. Zekiye Suludere Electron Microscopy Laboratory at the Gazi Uni-
versity, Faculty of Science. Terminology was adopted from Barthlott
(1981), Abid et al. (2016), Özbek & Uzunhisarcıklı,2020a;2020b, and
Abdel Khalik et al. (2021).
2.4 |Numerical analysis
The seed morphological characteristics of the 19 taxa, along with their
coefficients of correlation, were determined, and the taxa were
grouped using the hierarchical clustering analysis method.
Unweighted Pair Group Method with Arithmetic Mean (UPGMA) was
carried out with the Multivariate Statistical Package (MVSP) version
3.22 software (Kovach, 2013).
Principal component analysis (PCA) was performed to determine
the most significant characters that is, those accounting for the great-
est proportion of the variability. Thus, all the seed morphological char-
acters investigated were first subjected to this analysis, and the non-
significant variables showing a low percentage were eliminated.
Untransformed, centered, and unstandardized data were used to cre-
ate the correlation matrix. An eigenanalysis was then performed on
this matrix, and four eigenvectors were extracted. The eigenvalues
were plotted in a two-dimensional scatter plot of a Euclidean biplot
type to show both cases and variables along the first two principal
component axes (PC1, PC2) with the highest seed variation
(Özbek, 2022).
Eight seed morphological characters comprising two quantitative
(seed length and width) and six qualitative (color, seed surface orna-
mentations on dorsal and lateral sides, indumentum at dorsal and ven-
tral regions, and periclinal surface of epidermal cells) characters were
selected for the operational taxonomic units (OTUs) of the 19 taxa of
the genus Alcea (Table 2). The qualitative properties were coded in
the data matrix as (0, 1, 2, …), while the averages of the quantitative
features were also considered. A primary matrix was created using the
19 taxa (OTUs) and eight characters for the multivariate analysis.
Therefore, cluster analysis was conducted to group the 19 taxa into
clusters as a dendrogram based on the overall seed morphological var-
iable similarity. All computations were carried out using the MVSP
3.22 software.
3|RESULTS
The main seed macro- and micromorphological characteristics of the
Alcea taxa investigated were summarized in detail in Table 3. The vari-
ations in seed length and width for all taxa were shown in Figure 2.
LM and SEM micrographs were illustrated in Figures 3–9.
TABLE 2 Eight seed morphological characters to distinguish the 19 taxa of the genus Alcea L.
Taxa/characters
Seed
length
Seed
width Color
Ornamentation
(dorsal)
Ornamentation
(lateral)
Indumentum
(dorsal)
Indumentum
(ventral)
Periclinal
surface
Alcea acaulis 3.34 2.43 2 1 1 0 0 0
A. striata subsp.
striata
2.83 2.14 0 0 0 0 2 0
A. striata subsp.
rufescens
3.08 2.21 0 0 0 0 2 0
A. remotiflora 2.96 2.18 1 0 1 0 2 0
A. digitata 3.24 2.22 0 1 1 1 2 0
A. setosa 4 2.56 2 0 0 2 2 0
A. apterocarpa 3.4 2.29 0 0 1 1 2 0
A. kurdica 3.32 2.42 0 0 0 1 2 0
A. heldreichii 2.67 1.99 0 0 0 0 2 0
A. calvertii 2.39 1.93 0 0 0 0 1 0
A. hohenackeri 3.19 2.34 0 1 1 0 2 0
A. pisidica 3.43 2.15 0 2 1 1 2 1
A. guestii 3.75 2.11 0 1 0 1 2 0
A. biennis 2.69 2.18 3 0 1 0 0 0
A. dissecta 3.3 2.53 0 2 0 2 2 2
A. excubita 2.74 2.12 1 2 2 0 0 3
A. flavovirens 3.19 2.28 0 0 0 1 2 0
A. fasciculiflora 3.67 2.54 0 0 0 0 2 0
A. rosea 4.21 3.87 3 0 0 0 2 0
ÖZBEK and UZUNHISARCIKLI 5
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TABLE 3 Seed morphological data of Alcea L. (values in mm).
Species/characters Color
Length (L) Width (W)
L/W
Indumentum
Min. Max. Mean Min. Max. Mean Dorsal Ventral
Alcea acaulis Grayish-brown 2.92 3.7 3.34 2.07 2.89 2.43 1.37 ± 0.07 More sparsely More
sparsely
A. striata
subsp. striata
Dark brown 2.67 3 2.83 1.91 2.33 2.14 1.32 ± 0.06 More sparsely Densely
A. striata subsp.
rufescens
Dark brown 2.7 3.5 3.08 1.76 2.48 2.21 1.4 ± 0.11 More sparsely Densely
A. remotiflora Light to dark
brown
2.72 3.33 2.96 1.95 2.42 2.18 1.36 ± 0.07 More sparsely Densely
A. digitata Dark brown 3.03 3.43 3.24 1.99 2.56 2.22 1.46 ± 0.08 Sparsely Densely
A. setosa Grayish-brown 3.66 4.19 4 2.08 2.81 2.56 1.56 ± 0.1 Densely Densely
A. apterocarpa Dark brown 3.24 3.64 3.4 2.09 2.63 2.29 1.48 ± 0.1 Sparsely Densely
A. kurdica Dark brown 3.14 3.53 3.32 2.09 2.78 2.42 1.37 ± 0.08 Sparsely Densely
A. heldreichii Dark brown 2.48 2.85 2.67 1.76 2.24 1.99 1.34 ± 0.09 More sparsely Densely
A. calvertii Dark brown 2.22 2.72 2.39 1.83 2.11 1.93 1.23 ± 0.04 More sparsely Sparsely
A. hohenackeri Dark brown 2.64 3.48 3.19 2.05 2.6 2.34 1.36 ± 0.09 More sparsely Densely
A. pisidica Dark brown 2.93 4.19 3.43 1.88 2.59 2.15 1.6 ± 0.12 Sparsely Densely
A. guestii Dark brown 3.31 4.13 3.75 1.72 2.65 2.11 1.79 ± 0.16 Sparsely Densely
A. biennis Blackish-brown 2.57 2.82 2.69 2.05 2.43 2.18 1.23 ± 0.05 More sparsely More
sparsely
A. dissecta Dark brown 2.4 3.8 3.3 2.13 3.19 2.53 1.31 ± 0.15 Densely Densely
A. excubita Light to dark
brown
2.5 2.9 2.74 1.91 2.31 2.12 1.29 ± 0.06 More sparsely More
sparsely
A. flavovirens Dark brown 3 3.42 3.19 1.89 2.53 2.28 1.4 ± 0.08 Sparsely Densely
A. fasciculiflora Dark brown 3.38 3.92 3.67 2.3 2.73 2.54 1.44 ± 0.06 More sparsely Densely
A. rosea Blackish-brown 2.5 6.5 4.21 2 6.5 3.87 1.15 ± 0.17 More sparsely Densely
Species/
characters
Surface pattern Epidermal cell
shape
Anticlinal walls Periclinal walls
Dorsal Lateral Level Shape Level Surface
Alcea acaulis Reticulate-
rugulate
Reticulate-
rugulate
Polygonal Raised Straight or
undulate
Concave Striate or smooth
A. striata subsp.
striata
Reticulate Reticulate Polygonal or
rectangular
Raised or
flat
Straight or
undulate
Concave Striate or smooth
A. striata subsp.
rufescens
Reticulate Reticulate Polygonal or
rectangular
Raised or
flat
Straight or
undulate
Concave Striate or smooth
A. remotiflora Reticulate Reticulate-
rugulate
Polygonal Raised Straight or
undulate
Concave Striate or smooth
A. digitata Reticulate-
rugulate
Reticulate-
rugulate
Polygonal Raised Straight or
undulate
Concave Striate or smooth
A. setosa Reticulate Reticulate Polygonal or
rectangular
Raised or
flat
Straight Concave Striate or smooth
A. apterocarpa Reticulate Reticulate-
rugulate
Polygonal or
rectangular
Raised or
flat
Straight or
undulate
Concave Striate or smooth
A. kurdica Reticulate Reticulate Polygonal or
rectangular
Raised Straight Concave Striate or smooth
A. heldreichii Reticulate Reticulate Polygonal or
rectangular
Raised Straight or
undulate
Concave Striate or smooth
A. calvertii Reticulate Reticulate Polygonal or
rectangular
Raised or
flat
Straight or
undulate
Concave Striate or smooth
A. hohenackeri Reticulate-
rugulate
Reticulate-
rugulate
Polygonal or
rectangular
Raised Straight or
undulate
Concave Striate or smooth
6ÖZBEK and UZUNHISARCIKLI
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3.1 |Seed size, shape, and color
The seeds are strophiolate, reniform with a rounded apex and reni-
form base in shape. They show variation in color: light brown to dark
brown, grayish-brown, or blackish-brown. Their sizes ranges from
2.22 (Alcea calvertii) to 6.5 mm (A. rosea) in length, and 1.72 (A. guestii)
to 6.5 (A. rosea) mm in width. In all species studied, the hairs are white
and pilose on the ventral (especially around the hilum) and dorsal sides
of the seeds. The indumentum at these sides varies in density from
sparser, to sparse to dense. The hilum is found in the basal position.
3.2 |Seed surface
The seed surface is formed by polygonal or rectangular-shaped epi-
dermal cells. These cells consist of anticlinal walls showing a reticulate
appearance that is conspicuous or inconspicuous, straight or undulate,
and flat or raised, and periclinal walls with conspicuous, concave or
flat, and smooth, striate to ruminate surfaces. While most of the taxa
have polygonal and rectangular epidermal cells, only polygonal epider-
mal cells were found in three species: Alcea acaulis,A. remotiflora, and
A. digitata. The shape of the anticlinal walls is mostly straight or undu-
late, whereas Alcea setosa,A. kurdica, and A. dissecta have straight
anticlinal walls. These walls are generally distinct, but they are incon-
spicuous only in Alcea excubita. Two types of anticlinal walls of the
epidermal cells were observed, raised or flat. Raised and flat anticlinal
walls were dominant among the taxa being studied, but Alcea acaulis,
A. remotiflora,A. digitata,A. kurdica,A. heldreichii,A. hohenackeri, and
A. pisidica had only raised anticlinal walls. Periclinal walls are concave
except for Alcea excubita (concave or flat). Moreover, the surface pat-
terns of periclinal walls are generally striate or smooth in most species
investigated, in addition to striate or ruminate in Alcea pisidica, striate
to ruminate or smooth in A. dissecta, and the ruminate type occurs in
A. excubita.
TABLE 3 (Continued)
Species/
characters
Surface pattern Epidermal cell
shape
Anticlinal walls Periclinal walls
Dorsal Lateral Level Shape Level Surface
A. pisidica Reticulate-
ruminate
Reticulate-
rugulate
Polygonal or
rectangular
Raised Straight or
undulate
Concave Striate or ruminate
A. guestii Reticulate-
rugulate
Reticulate Polygonal or
rectangular
Raised or
flat
Straight or
undulate
Concave Striate or smooth
A. biennis Reticulate Reticulate-
rugulate
Polygonal or
rectangular
Raised or
flat
Straight or
undulate
Concave Striate or smooth
A. dissecta Reticulate-
ruminate
Reticulate Polygonal or
rectangular
Raised or
flat
Straight Concave Striate to ruminate or
smooth
A. excubita Reticulate-
ruminate
Reticulate-
ruminate
Polygonal or
rectangular
Raised or
flat
Straight or
undulate
Concave or
flat
Ruminate
A. flavovirens Reticulate Reticulate Polygonal or
rectangular
Raised or
flat
Straight or
undulate
Concave Striate or smooth
A. fasciculiflora Reticulate Reticulate Polygonal or
rectangular
Raised or
flat
Straight or
undulate
Concave Striate or smooth
A. rosea Reticulate Reticulate Polygonal or
rectangular
Raised or
flat
Straight or
undulate
Concave Striate or smooth
FIGURE 2 Box-plot graph for seed length and width in the studied Alcea L. taxa. The horizontal bar inside the rectangle is the median, the
rectangle shows 50% of the interquartile distance, the ends show the amplitude variation, the circles correspond to the outliers, and the stars
indicate the extremes.
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All taxa have small wartlets showing verrucate ornamentations on
the seeds, and their seed coat patterns vary from reticulate,
reticulate-rugulate to reticulate-ruminate. There are some differences
in seed surface ornamentations among the taxa studied, and also
between the dorsal and lateral sides of the seeds of the taxa. Alcea
striata subsp. striata,A. striata subsp. rufescens,A. setosa,A. kurdica,
A. heldreichii,A. calvertii,A. flavovirens,A. fasciculiflora, and A. rosea
show reticulate ornamentations on both dorsal and lateral regions.
Similarly, reticulate-rugulate surface patterns on two faces have been
observed in Alcea acaulis,A. digitata, and A. hohenackeri. The surface
sculpturing on the dorsal sides is reticulate, whereas it is reticulate-
rugulate on the lateral sides in Alcea remotiflora,A. apterocarpa, and
A. biennis. While Alcea pisidica,A. dissecta, and A. excubita show
reticulate-ruminate ornamentations on the dorsal sides, it was found
to be reticulate-rugulate, reticulate, and reticulate-ruminate on the lat-
eral sides, respectively. Finally, the surface sculpturing is reticulate-
rugulate on the dorsal regions, and reticulate on the lateral regions in
Alcea guestii.
FIGURE 4 Light microscope
micrographs of seeds of Alcea
guestii (1), A. biennis (2), A. dissecta
(3), A. excubita (4), A. flavovirens
(5), A. fasciculiflora (6), A. rosea (7).
FIGURE 3 Light microscope
micrographs of seeds of Alcea
acaulis (1), A. striata subsp. striata
(2), A. striata subsp. rufescens (3),
A. remotiflora (4), A. digitata (5),
A. setosa (6), A. apterocarpa (7),
A. kurdica (8), A. heldreichii (9),
A. calvertii (10), A. hohenackeri
(11), A. pisidica (12).
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3.3 |Numerical analysis
The phenetic relationships of the Alcea taxa reflected in the seed
macro- and micromorphological variations were presented using two
statistical analyses. First, PCA was performed to assess which vari-
ables were important in explaining the total variation among the eight
cases investigated. The values obtained for the eigenvectors and the
total cumulative variance were presented in Table 4, and the scatter
plot with the seed morphological characteristics were shown in
Figure 10. The PCA of the eight variables explains 90.761% of the
observed variation with the first four axes. The first principal compo-
nent (PC1) explains 35.629% of the total variation in the taxa studied.
The color, indumentum at the ventral region, and seed coat ornamen-
tations on the lateral sides are the most significant variables in the
first principal component because they have the highest relative varia-
tion rate. The periclinal surface of epidermal cells, seed coat ornamen-
tations on the dorsal region, and color have the strongest influence on
the taxa in the second principal component (PC2), which explains
32.242% of the variance. The third principal component (PC3)
accounted for 17.348% of the variation and is associated with the
indumentum at the dorsal side, with seed length and color being the
most significant variables. The fourth principal component (PC4)
explains 5.541% of the variance, mainly through variables such as
indumentum at the dorsal and ventral regions and seed width
(Table 4).
A dendrogram was constructed using cluster analysis based on
the eight seed morphological characteristics (seed length and width,
color, seed coat ornamentations on dorsal and lateral sides, indumen-
tum at dorsal and ventral faces of seed, and periclinal surface of epi-
dermal cells) for the OTUs of the 19 taxa (Figure 11). The cluster
analysis divides the taxa into four main groups, namely, A, B, C, and D,
with 62.2% similarity. Alcea dissecta is distinctly separated from the
other taxa in the cluster A with 63.3% similarity. Cluster B consists of
Alcea rosea and A. setosa. Cluster C includes two subgroups, namely,
C1 and C2. In the first subgroup C1, three of the four taxa are mor-
phologically similar (Alcea pisidica,A. guestii, and A. hohenackeri,
83.1%). Cluster C2 is divided into subgroups C2a and C2b, with C2a
consisting of Alcea calvertii, with 82.1% similarity. Cluster C2b
FIGURE 5 Scanning electron microscope micrographs of seeds of Alcea acaulis (1–4), A. striata subsp. striata (5–8), A. striata subsp. rufescens
(9–12), A. remotiflora (13–16).
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comprises Alcea remotiflora,A. flavovirens,A. kurdica,A. apterocarpa,
A. fasciculiflora,A. striata subsp. rufescens,A. heldreichii, and A. striata
subsp. striata.Alcea flavovirens and the morphologically close
A. kurdica are quite similar (98.2%). Alcea heldreichii,A. striata subsp.
striata and the other subspecies A. striata subsp. rufescens are mor-
phologically related with 96.8% similarity. Alcea excubita falls within
cluster D, with 62.2% similarity. Cluster D also comprises Alcea biennis
and A. acaulis.
Taxonomic key for Turkish Alcea based on macro-
micromorphological characters of seed
1+Seeds densely hairy at dorsal side 2
1Seeds sparsely or more sparsely hairy at
dorsal side
3
2+Seed coat ornamentation reticulate on
dorsal surface
A. setosa
2Seed coat ornamentation reticulate-
ruminate on dorsal surface
A. dissecta
3+Epidermal cell shape polygonal 4
3Epidermal cell shape polygonal or
rectangular
6
4+Seeds light to dark brown or dark brown 5
4Seeds grayish-brown A. acaulis
5+Seed coat ornamentation reticulate on
dorsal surface
A. remotiflora
5Seed coat ornamentation reticulate-
rugulate on dorsal surface
A. digitata
6+Periclinal walls concave 7
6Periclinal walls concave or flat A. excubita
7+Seed coat ornamentation reticulate or
reticulate-rugulate on dorsal surface
8
7Seed coat ornamentation reticulate-
ruminate on dorsal surface
A. pisidica
8+Anticlinal walls shape straight or undulate 9
8Anticlinal walls shape straight A. kurdica
9+Anticlinal walls raised 10
9Anticlinal walls raised or flat 11
10+Seed coat ornamentation reticulate A. heldreichii
10Seed coat ornamentation reticulate-
rugulate
A. hohenackeri
11+Seed coat ornamentation reticulate on
dorsal surface
12
11Seed coat ornamentation reticulate-
rugulate on dorsal surface
A. guestii
12+Seeds sparsely or more sparsely hairy at
ventral side
13
12Seeds densely hairy at ventral side 14
13+Seed coat ornamentation reticulate on
lateral surface
A. calvertii
13Seed coat ornamentation reticulate-
rugulate on lateral surface
A. biennis
14+Seeds mean length less than 3.67 mm, dark
brown
15
14Seeds mean length more than 3.67 mm,
blackish-brown
A. rosea
15+Seed coat ornamentation reticulate on
lateral surface
16
15Seed coat ornamentation reticulate-
rugulate on lateral surface
A. apterocarpa
16+Seeds mean length less than 3.19 mm 17
16Seeds mean length more than 3.19 mm A. fasciculiflora
17+Seeds more sparsely hairy at dorsal side A. striata
17Seeds sparsely hairy at dorsal side A. flavovirens
4|DISCUSSION
The seed morphological characteristics of the family Malvaceae are
consistent and very useful for the delimitation of taxa in this family
(Abid et al., 2016; Ahmed & Qaiser, 1989; Ather et al., 2009; Kirkbride
et al., 2006; Özbek & Uzunhisarcıklı,2020a;2020b). SEM has
revealed the significant morphological features for the identification
of species (Bona, 2020; Luqman et al., 2019; Özbek et al., 2018;
Özbek & Uzunhisarcıklı,2020a;2020b). In this research, LM and SEM
were used to observe the seed macro- and micromorphological prop-
erties for members of Alcea from Turkey. Patil et al. (2015) reported
that seed dimensions show great variation among the Abelmoschus
species within the Malvaceae family. The sizes of seeds were variable
among the taxa investigated, ranging from 2.22 to 6.5 mm in length
and 1.72 to 6.5 mm in width. The smallest seeds were observed in
Alcea calvertii with L: 2.39 ± 0.11 mm, W: 1.93 ± 0.08 mm, while the
largest ones were noted in A. rosea with L: 4.21 ± 1.55 mm, W: 3.87
± 1.83 mm. Alcea rosea as the cultivator form showed a wide variation
in their seed sizes. According to the data by Abid et al. (2016), the
seed sizes of Alcea rosea (L: 2.5 mm, W: 3.6 mm) and A. biennis (men-
tioned as A. pallida (Willd.) Waldst. & Kit.) (L: 2 mm, W: 2.5 mm) were
mostly similar to the findings in this study (L: 4.21 mm, W: 3.87 mm,
and L: 2.69 mm, W: 2.18 mm, respectively), whereas the results of
Bojˇ
nanský and Fargaˇ
sová (2007) for Alcea rosea (3.3–3.7 2.7–
3 mm) and A. biennis (mentioned as A. pallida) (2.8–3.2 4.2–4.6 mm)
did not match the data of our study.
Duke (1961), Ahmed and Qaiser (1989) and Abdel Khalik et al.
(2021) stated that seed color is another good reliable constant charac-
ter, and is widely used for the delimitation of various taxa. Most of
the taxa studied have dark brown seeds, and Alcea remotiflora and
A. excubita showed some degree of variation from light to dark.
Grayish-brown was recognized in the seeds of Alcea acaulis and
A. setosa. However, the seeds are blackish-brown in both taxa of Alcea
biennis as well as A. rosea. The seed color of Alcea rosea determined in
this study does not match the findings (grayish-brown) of Bojˇ
nanský
and Fargaˇ
sová (2007), the results (grayish black) of Thakor (2009) and
the (reddish brown) of Abid et al. (2016). In the study by Abid et al.
(2016), Alcea biennis was found to be dusty brown, different from the
current research.
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Esteves (2004) found no significant differences in the Pavonia
species being studied for seed shape. The researcher observed the
reniform seeds in most species being studied except for sect. Astero-
chlaena Garcke and sometimes in sect. Lopimia Mart. (obovoid shape).
Similarly, Özbek & Uzunhisarcıklı(2020a;2020b) stated that Turkish
Althaea and Lavatera L. species have reniform-shaped seeds. The seed
shape is rather uniform in the Alcea taxa studied. They are reniform
with a rounded apex and base reniform, strophiolate, and show hilum
at basal in agreement with Paul and Nayar (1987) for Alcea rosea, and
Bojˇ
nanský and Fargaˇ
sová (2007) for A. biennis and A. rosea. Thakor
(2009) stated that Alcea rosea has sub-reniform seeds. Abid et al.
(2016) reported that the seed shape in Alcea biennis and A. rosea was
reniform-transversally elliptic pyriform with a rounded apex and
reniform base.
El Naggar (2001) demonstrated the utility of hair characteristics
for distinguishing the genus Abelmoschus from Hibiscus. Patil et al.
(2020) detected that seed trichome features would play an important
role in taxonomy of the family Malvaceae. They also observed a con-
siderable variation in the trichome distribution on the seed surface
among Abelmoschus and Hibiscus taxa. Similarly, the indumentum
shows differences among the taxa studied for their distribution with
density in this research. The hairs are densely pilose on the ventral
sides (especially around the hilum) and more sparsely pilose on the
dorsal sides of the seeds in general. The seeds of Alcea digitata,
A. apterocarpa,A. kurdica,A. pisidica,A. guestii, and A. flavovirens are
sparsely hairy on the dorsal side, whereas A. setosa and A. dissecta
have dense hairs on this side. The remaining taxa indicate sparser
hairy on this region of the seeds. However, Alcea acaulis,A. biennis,
and A. excubita showed sparser hairy on the ventral regions of the
seeds, and only A. calvertii had sparse hairs. The other taxa except
these species mentioned are densely hairy in this region. Abid et al.
(2016) stated that the seed indumentum for Alcea biennis was sparsely
pubescent (puberulose) and was hairy upperside (tomentose) for
A. rosea, whereas Bojˇ
nanský and Fargaˇ
sová (2007) found this feature
to be glabrous for these two Alcea species. Their results were differ-
ent from the current study with pilose sparser hairy for Alcea biennis
and pilose sparser hairy on the dorsal sides and densely hairy on the
ventral ones for A. rosea. The hilum with some appressed hairs was
FIGURE 6 Scanning electron microscope micrographs of seeds of Alcea digitata (1–4), A. setosa (5–8), A. apterocarpa (9–12), A. kurdica
(13–16).
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observed in Alcea rosea by Paul and Nayar (1987) and were partly like
the results of this research.
Paul and Nayar (1987), Sivarajan and Pradeep (1996), Esteves
(2004), El Naggar (2001), and Abdel Khalik et al. (2021) reported that
the seed coat sculpturing patterns play a significant role in the differ-
entation of the Malvaceae members. Presently, considerable varia-
tions in the seed coat patterns were observed between the dorsal and
ventral regions and among the taxa. The Alcea taxa investigated have
small wartlets on their seeds, and so the Alcea seeds are verrucate.
However, they mostly show reticulate ornamentations on their seed
surfaces. The seeds of Alcea acaulis,A. digitata, and A. hohenackeri
have reticulate-rugulate ornamentations on the dorsal and ventral sur-
faces, while A. excubita showed reticulate-ruminate patterns on these
sides. Furthermore, Alcea remotiflora,A. apterocarpa,A. pisidica,
A. guestii,A. biennis, and A. dissecta demonstrated different ornamen-
tations between the dorsal and ventral regions of the seeds. Alcea
remotiflora,A. apterocarpa,A. pisidica, and A. biennis have reticulate-
rugulate ornamentations on the lateral faces of their seeds, while the
dorsal sides showed reticulate patterns in A. remotiflora,
A. apterocarpa, and A. biennis, and reticulate-ruminate in A. pisidica.
Similarly, Alcea guestii and A. dissecta showed reticulate ornamenta-
tions on the lateral sides of the seeds, but the seed coat patterns were
reticulate-rugulate in A. guestii on the dorsal region, and reticulate-
ruminate in A. dissecta. Pakvaran and Ghahreman (2005) found simple
reticulate patterns in Alcea mazandaranica Pakravan & Ghahr. and
A. iranshahrii Pakravan, Ghahr. & Assadi. According to the data by
Abid et al. (2016), the seed surface patterns of Alcea biennis and
A. rosea (reticulate-foveate, verrucate) are mostly congruent with
those of A. biennis (verrucate, reticulate on dorsal sides and reticulate-
rugulate on ventral ones) and A. rosea (verrucate, reticulate) in the pre-
sent research. The seed coat sculpturing found in this study for Alcea
rosea (reticulate) agrees with that of Paul and Nayar (1987)
(reticulate).
The epidermis cell shapes of the seed coats are generally polygo-
nal or rectangular in most species investigated, but an only polygonal
type occurred in Alcea acaulis,A. remotiflora, and A. digitata. Barthlott
(1981) detected that the anticlinal and periclinal walls of the epidermis
cells have high taxonomic values at interspecific levels in different
FIGURE 7 Scanning electron microscope micrographs of seeds of Alcea heldreichii (1–4), A. calvertii (5–8), A. hohenackeri (9–12), A. pisidica
(13–16).
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families. Our findings are congruent with Barthlott (1981) only regard-
ing the ornamentations of the periclinal walls. The surfaces of the
periclinal walls are striate or smooth in most of the taxa, whereas stri-
ate or ruminate types have been observed in Alcea pisidica, striate to
ruminate or smooth in A. dissecta, and ruminate in A. excubita. No sig-
nificant differences in anticlinal and periclinal wall structures were
observed among the Alcea species investigated. The anticlinal walls
are conspicuous or inconspicuous, straight or undulate, or only
straight, flat or raised or only raised, and the periclinal walls are con-
spicuous, and concave or concave or flat in all taxa examined. The sur-
face of the periclinal walls were found to be smooth in Alcea rosea by
El Naggar (2001) and are partly similar to our study (striate or smooth),
while El Naggar's observations about the periclinal walls being flat for
this species differed from our study (concave).
The PCA was performed to determine which variables were signif-
icant in explaining the total variation among the 19 taxa investigated.
This analysis showed that the seed length and width, color, seed coat
ornamentations on the dorsal and lateral sides, indumentum at the dor-
sal and ventral faces of the seed, and periclinal surface of epidermal
cells are the most important properties with the highest relative
variation rate. The PCA and UPGMA based on eight seed morphologi-
cal character states indicate that the Turkish Alcea taxa are divided into
four main groups. Alcea dissecta is separated from the other species at
first with reticulate-ruminate ornamentations on the dorsal sides, retic-
ulate patterns on the ventral faces, and striate to ruminate or smooth
periclinal walls of seeds among the species studied in Cluster A. Alcea
rosea and A. setosa display a 73% similarity with the largest seeds in
Cluster B. Alcea pisidica,A. guestii, and A. hohenackeri are placed in
Cluster C1. The similarity rate among these species is 83% given their
close measurements in seed size, seed color, and indumentum. They
are also morphologically similar species because they both possess
mature mericarps with prominent wings and an epicalyx more than ½
as long as sepals or equal. The closest of these three species based on
seed characteristics in the dendrogram strongly supports their mor-
phological resemblance. In Cluster C2, Alcea flavovirens has 98.2% simi-
larity with A. kurdica for their seed color, indumentum at dorsal and
ventral sides, seed coat ornamentations on the dorsal and ventral faces
and epidermal cells' periclinal surface patterns. These findings are con-
gruent with the morphological resemblance of Alcea flavovirens and
A. kurdica with lobed leaves, epicalyx less than ½ as long as the calyx,
FIGURE 8 Scanning electron microscope micrographs of seeds of Alcea guestii (1–4), A. biennis (5–8), A. dissecta (9–12), A. excubita (13–16).
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petal length nearly 40 mm and mericarps being distinctly winged. Alcea
striata subsp. rufescens, the other subspecies A. striata subsp. striata,
and A. heldreichii placed ongoing in this cluster have undivided or
slightly lobed leaves and epicalyx less than ½ as long as the calyx. The
similarity rate among these allied taxa is >96% according to the seed
morphological findings. They are similar based on their seed color,
indumentum at the dorsal and ventral regions, seed surface sculpturing
on the dorsal and ventral sides, and periclinal surface of epidermal
cells. The results according to the cluster analysis based on the seed
macro- and micromorphology are partly in accordance with the sys-
tematics based on the macromorphological properties of the Alcea taxa
investigated.
5|CONCLUSION
The seeds of Alcea members vary in their size, color, indumentum,
and surface pattern features. Cluster analysis revealed four main
groups based on both qualitative and quantitative data, and its find-
ings are partially correlate with the classification of the species in
this genus. PCA showed that color, indumentum at the ventral
region, seed coat ornamentations on dorsal and lateral sides, and
periclinal surface of epidermal cells were the most valuable charac-
teristics in explaining the total variation among the taxa examined.
According to the PCA and cluster analyses based on the seed mor-
phological characteristics of 19 Alcea taxa, seed length and width,
FIGURE 9 Scanning electron microscope micrographs of seeds of Alcea flavovirens (1–4), A. fasciculiflora (5–8), A. rosea (9–12).
TABLE 4 Principal component
analysis variable loadings for the first
four components.
Variables PC 1 PC 2 PC 3 PC4
Seed length (L) 0.028 0.102 0.447 0.283
Seed width (W) 0.082 0.145 0.302 0.424
Color 0.724 0.424 0.353 0.030
Seed coat ornamentation (dorsal) 0.141 0.583 0.215 0.208
Seed coat ornamentation (lateral) 0.293 0.249 0.177 0.142
Indumentum (dorsal) 0.182 0.163 0.619 0.683
Indumentum (ventral) 0.524 0.097 0.319 0.440
Periclinal surface of epidermal cells 0.235 0.593 0.147 0.127
Eigenvalues 1.507 1.363 0.734 0.234
Percentage 35.629 32.242 17.348 5.541
Cumulative percentage 35.629 67.871 85.220 90.761
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seed color, seed coat ornamentations, indumentum, and periclinal
surface pattern of epidermal cells prove to be important morpho-
logical characteristics for the discrimination at a specific rank. The
present study clarifies the significance of LM and SEM for the cor-
rect and detailed identification, and species separation of the genus
Alcea based on seed features. This research provides comprehen-
sive seed morphological knowledge on the taxonomic aspects of
the genus Alcea. Moreover, this work can be useful for the seed
morphological investigations of other related genera within
Malvaceae.
AUTHOR CONTRIBUTIONS
Funda Özbek: Investigation; writing –review and editing;
writing –original draft; methodology; conceptualization. Mehmet
Erkan Uzunhisarcıklı:Investigation; funding acquisition; writing –
original draft.
FIGURE 10 Scatter plot of the principal component analysis (PCA) for seed morphological characteristics in Alcea L. taxa along PC1 and PC2.
V1, seed length; V2, seed width; V3, color; V4, seed coat ornamentation (dorsal); V5, seed coat ornamentation (lateral); V6, indumentum (dorsal);
V7, indumentum (ventral); V8, periclinal surface of epidermal cells.
FIGURE 11 Cluster dendrogram of the investigated Alcea L. taxa based on seed morphological data.
ÖZBEK and UZUNHISARCIKLI 15
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ACKNOWLEDGMENTS
We are grateful to thank to the Scientific and Technological Research
Council of Turkey for the collection of plant materials (TUBITAK-
TBAG 2282) for financial support, and also thank to the curators of
herbaria GAZI who allowed us to study their Alcea material.
CONFLICT OF INTEREST STATEMENT
The authors declare no conflict of interest.
DATA AVAILABILITY STATEMENT
The data that support the findings of this study are available from the
corresponding author upon reasonable request.
ORCID
Funda Özbek https://orcid.org/0000-0002-0135-0155
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How to cite this article: Özbek, F., & Uzunhisarcıklı,M.E.
(2023). Taxonomic significance of seed
macro-micromorphology of Turkish Alcea L. (Malvaceae)
through light microscopy and scanning electron microscopy.
Microscopy Research and Technique,1–17. https://doi.org/10.
1002/jemt.24385
ÖZBEK and UZUNHISARCIKLI 17
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