Bangladesh J. Plant Taxon. 30(2): 201-212, 2023 (December) DOI: https://doi.org/10.3329/bjpt.v30i2.70497 © 2023 Bangladesh Association of Plant Taxonomists ANATOMY, POLLEN AND SEED MORPHOLOGY OF ENDEMIC SPECIES VERBASCUM GLOBIFERUM HUB.-MOR. AND V. LYSIOSEPALUM HUB.-MOR. (SCROPHULARIACEAE) IN DIYARBAKIR, TURKEY AND THEIR TAXONOMIC IMPORTANCE MURAT KILIÇ* Department of Crops and Animal Production, Mardin Artuklu University, 47200 Mardin, Artuklu, Turkey Keywords: Anatomy; Diyarbakır; Pollen morphology; Seed surface; SEM; Verbascum. Abstract Verbascum globiferum Hub.-Mor. and V. lysiosepalum Hub.-Mor. are endemic to Turkey which is a center of endemism for Verbascum species. This study gives the anatomical, palynological and seed micromorphological features of V. globiferum and V. lysiosepalum which grow in Diyarbakır. The xylem elements occupy a large area in the root cross-sections of the species. In stem cross-sections, the upper part of the epidermis cells is surrounded by a separate cuticle layer and the pith region covers a large area. In the leaves, the main vein is shapped as collateral bundle. Pollen grains are tricolporate- tricolpate, oblate-spheroidal, and exine ornamentation is reticulate. Seeds are brown and oblong-ovate to prismatic and alveolate. The seed coat ornamentation is irregular with polygonal cells, with densely, distinct vesicles.The capsules are pubescent, covered with stellate and branched hairy or glandular hairs.The seed displayed that substantial taxonomic understanding can be acquired from examining the seed characteristics of Verbascum, particularly at the species level. Intrudiction Verbascum L. genus, which is also known as "Sığırkuyruğu" in Anatolia, is one of the largest genera of the Scrophulariaceae family (Heywood, 1993). The genus Verbascum with 360 species (Judd et al., 1999) worldwide and 257 species and 132 supplement hybrids in Turkey, and has been divided into 13 artificial groups in Turkey (Huber-Morath, 1978; Davis et al., 1988; Karavelioğulları, 2012).The endemism ratio of the genus is very high, with 202 endemic species (80%). (Huber-Morath, 1978; Davis et al., 1988; Karavelioğulları, 2012; Karaveliogulları et al., 2014; Firat, 2022). The genus Verbascum is one of the largest genera in terms of the number of species it contains, which is known to have problems in diagnosis and taxonomy because it shows a lot of hybridization in general. There are few studies on the morphological and anatomical features of the genus (Çakır and Bağcı, 2006). There are many SEM studies based on pollen morphology of the some Verbascum species taxa (Juan et al., 1997; Dane and Yilmaz, 2002; Kheyri, 2009; Asmat et al., 2011; Al-Hadeethy et al., 2014; Öztürk et al., 2018; Aktas, 2019; Aktas et al., 2020; Baser, 2021). Seed micromorphology of the genus Verbascum has been examined by several researchers including Juan et al. (1997), Petkovié et al. (1997), Attar et al. (2007), Kheiri et al. (2009), Cabi et al. (2011), Aktas 2019, and Aktas et al. 2020. Anatomical studies of this genus were made by few researchers (Özdemir and Altan, 2007; Kheiri et al., 2009; Yılmaz and Dane, 2011; Alan and Gökman, 2015; Küçük, 2017; Tekin and Yılmaz, 2018; Aktas, 2019; Aktaset al., 2020; Küçük et al., 2021). There are still deficiencies in information and many taxa are not studied yet in Turkey. * E-mail: muratkilic04@gmail.com 202 KILIÇ Verbascum globiferum and V. lysiosepalum are species that grow in Diyarbakır in the Irano- Turanian phytogeographic region and are endemic to Turkey (Huber-Morath, 1978). In this study, the root, stem and leaf anatomy, pollen, and seed micromorphology of the endemic V. globiferum and V. lysiosepalum species have been investigated for the first time to asses their taxonomic values. Material and Methods The specimens belonging to Verbascum globiferum and V. lysiosepalum were collected from Ergani and between Siverek and Diyarbakır (within the borders of Diyarbakır Province) localities in Diyarbakır (Fig. 1). We have deposited the voucher samples in the herbarium of the Department of Plants and Animal Production of Kızıltepe Vocational School, Mardin Artuklu University (Collector numbers; M.Kılıç 351, M.Kılıç 395, M.Kılıç 353, M.Kılıç 355, M.Kılıç 356, M.Kılıç 357-1, M.Kılıç 394) . The taxonomic description of the plant was prepared according to Davis et al. (1988) and Karavelioğulları (2012). Fig. 1. Distribution map of Verbascum globiferum (*) and V. lysiosepalum (°). Collected specimens were preserved in falcon tubes in 70% alcohol for use in anatomical studies. Sections taken from the root, stem and leaf parts of the plants with the help of a razor were prepared by staining with safranin-fast green and examined under the light microscope (4x and 10x) and photographed (Bozdağ et al., 2016). For palynological examinations, light (LM) and (SEM) values of all pollen grains were determined by standard methods described by Erdtman (1952). Pollen grains for LM examination were prepared following the standard procedure of Wodehouse (1935). Thirty pollen grains per specimen were regarded as sufficient for the palynological analysis (Wodehouse, 1935; Kheiri et al., 2006; Cabi et al., 2011). For SEM, pollen were removed by distilled water treatment, the air-dried, pollens were directly mounted on stubs using double-sided adhesive tape and it was covered with gold. The photomicrographs were taken with a ZEISS EVO 50 scanning electron microscope. The values of P (polar axis length), E (equatorial diameter), Clg (Colpus longitude), Clt (Colpus latitude), Plg (Polar longitude), Plt (Polar latitude), Ex (Exine thickness), and In (Intine thickness) were measured, and the P/E ratio was calculated, Apt (Aperture type), and Or (Ornamentation) for 30 pollen grains were measured under light microscope.. The terminology of the pollen follows that of Punt et al. (2007). The values are presented as minimum, maximum and mean, that is represented in Table 2. ANATOMY, POLLEN AND SEED MORPHOLOGY OF ENDEMIC SPECIES 203 Seeds were first examined using a Isolab stereomicroscope to ensure that they were of normal size and mature. In order to determine the average seed sizes, 30 mature seeds were measured. For SEM, seed debris were removed by distilled water treatment, the air-dried seeds were then mounted on stubs and it was covered. The photomicrographs were taken with ZEISS EVO 50 scanning electron microscope. Terminology for descriptions of morphological characteristics of the mericarps were followd by Sutton (1988), Juan et al. (1997), Attar et al. (2007) and Cabi et al. (2011). Results and Discussion In this study, various features of root, stem and leaf anatomical structures, pollen, and seed surface structures of endemic Verbascum globiferum and V. lysiosepalum species are stated. Biometric measurements of the root, stem, and leaf tissues and cells are given in Table 1 and shown in Figs 2, 3, 4. The characteristics of pollen grains are summarized in Table 2 and shown in Fig. 5. The morphological characteristics of the seed grains, including their size, shape, color, and surface characteristics, are summarized in Table 3 and shown in Fig. 6. The morphological characteristics of the capsule grains, including their size, shape, and color characteristics, are summarized in Table 4 and shown in Fig. 7. Anatomy Root anatomy: Cross-sections taken from the root of Verbascum globiferum have revealed that the periderm layer on the outermost surface of the root is thin and its cells are irregular. A multilayered parenchyma is present under the periderm. Below the parenchyma are 3-5 layered phloem cells. The cambium is indeterminate. The xylem covers a larger area and fills the middle of the root. Trachea cells are irregularly located, larger than the tracheit cells, and their length is greater than their width. Phloem occupies a narrower area than the xylem. Pith rays comprise 2-6 rowed rectangular cells. The pith consists of polygonal or orbicular parenchymatous cells (Fig. 2, Table 1). Fig. 2. Cross-section of the root of A: V. globiferum, B: V. lysiosepalum. Pe: Periderm, P: Parenchyma, Ph: Phloem, X: Xylem, Pr: Pith ray, Tr: Trachea, Pt: Pith region. 204 KILIÇ Table 1. The anatomical mesurements of Verbascum globiferum and V. lysiosepalum. Width (µ) Length(µ) Species / Tissues Min. Mak. Mean±S. Min. Mak. Mean±S. V. globiferum Root Peridermis cell 12.20 51.16 27.53±10.53 9.75 31.93 17.83±6.53 Parenchyma cell 10.93 63.18 36.15±17.82 6.34 30.82 17.87±7.99 Phloem cell 6.63 24.27 15.53±5.20 3.21 14.15 7.02±2.69 Trachea cell 20.04 52.17 35.14±8.17 16.35 59.08 36.44±12.81 V. lysiosepalum Peridermis cell Parenchyma cell Phloem cell Trachea cell 7.76 13.13 9.44 29.41 38.50 75.65 19.78 96.77 19.58±7.61 36.08±17.64 12.90±2.85 58.41±22.72 7.74 7.20 24.16 27.40 14.48±4.40 17.07±6.36 6.03 27.46 15.66 81.70 10.76±2.72 55.26±16.88 V. globiferum Stem Cuticle - - - 6.61 14.00 9.91±1.95 Epidermis cell 10.15 32.49 18.68±5.86 9.25 24.64 15.18±4.97 Collenchyma cell 6.82 43.91 18.36±11.68 7.41 48.06 17.92±11.25 Parenchyma cell 15.21 48.36 27.46±8.99 9.14 24.00 17.04±4.90 Phloem cell 4.50 18.17 9.58±3.20 3.26 6.60 4.92±0.96 Trachea cell 10.57 26.91 18.08±4.81 11.54 27.42 20.41±3.98 Pith cell 28.60 159.25 96.98±42.62 24.08 189.62 93.30±51.01 V. lysiosepalum Cuticle Epidermis cell Collenchyma cell Parenchyma cell Phloem cell Trachea cell Pith cell - 11.17 6.51 14.27 3.15 13.72 31.10 - 28,28 20.13 35.27 11.47 32.38 106.74 - 17.77±3.97 12.04±4.00 24.21±5.51 7.26±2.27 23.30±6.33 64.65±23.83 3.55 8.89 7.64 9.87 2.60 8.63 29.39 10.40 20.62 18.21 27.67 9.68 45.20 103.32 7.09±1.95 13.33±3.17 12.41±3.02 18.52±4.40 5.79±2.24 27.50±10.33 64.49±24.51 V. globiferum Leaf Cuticle - - - 5.15 13.90 8.57±2.36 Upper epidermis cell 11.20 70.26 34.40±12.71 8.24 27.30 18.41±5.41 Palisade parenchyma 13.09 22.29 16.83±2.61 26.00 48.52 33.51±5.89 Spongy parenchyma 11.73 22.78 16.92±2.82 12.33 27.00 20.73±5.00 Mesophyll layer - - - 174.55 332.09 238.22±36.68 Lower epidermis cell 7.40 22.90 14.96±4.97 9.25 20.39 13.52±2.71 V. lysiosepalum Cuticle - - - 4.65 13.32 8.28±2.09 Upper epidermis cell 11.39 25.34 18.21±4.24 8.76 30.04 16.79±5.00 Palisade parenchyma 7.77 17.36 11.06±2.41 22.22 44.03 31.21±5.00 Spongy parenchyma 8.85 22.49 14.88±3.91 8.28 30.86 19.10±6.93 Mesophyll layer - - - 185.64 237.59 210.58±15.21 Lower epidermis cell 8.99 21.84 14.08±3.56 7.24 14.10 11.08±1.75 ANATOMY, POLLEN AND SEED MORPHOLOGY OF ENDEMIC SPECIES 205 Cross-sections taken from the root of V. lysiosepalum have revealed that the periderm layer on the outermost surface of the root is thin and its cells are irregular. A multilayered parenchyma is present under the periderm. Below the parenchyma are 2-4 layered phloem cells. The cambium is indeterminate. The xylem covers a larger area and fills the middle of the root. Trachea cells are irregularly located, larger than the tracheit cells, and their width is greater than their length. Phloem occupies a narrower area than the xylem. Pith rays comprise 2-6-rowed rectangular cells. The pith consists of polygonal or orbicular parenchymatous cells (Fig. 2, Table 1). Stem anatomy: Cross-sections taken from the stem of V. globiferum have exhibited a monolayer epidermis covered by an undulate cuticle. The epidermis is composed of oval or rectangular cells. There are glandular and eglandular hairs on the epidermis. Underneath the epidermis, there are 3-5 layers of collenchyma cells. The parenchyma tissue consists of 7-10 layers of oval and orbicular parenchymatous cells. Under the parenchyma, there are 4-6 layers of sclerenchyma. Cambium is indistinguishable. The xylem occupies a larger area than phloem. The pith comprises hexagonal or orbicular parenchymatous cells with intercellular spaces (Fig. 3, Table 1). Fig. 3. Cross-section of the stem of A: V. globiferum, B: V. lysiosepalum. Eg: Eglandular hair, Ch: Compound hair, Cu: Cuticle, Ep: Epidermis, Co: Collenchyma, P: Parenchyma, Sc: Sclerenchyma, Ph: Phloem, X: Xylem, Tr: Trachea, Pt: Pith region. Cross-sections taken from the stem of V. lysiosepalum have exhibited a monolayer epidermis covered by an undulate cuticle. The epidermis is composed of oval, ovate or rectangular cells. There are compound, glandular, and eglandular hairs on the epidermis. Underneath the epidermis, there are 3-5 layers of collenchyma cells. The parenchyma tissue consists of 7-10 layers of oval and orbicular parenchymatous cells. Under the parenchyma, there are 5-7 layers of sclerenchyma. Cambium is indistinguishable. The xylem occupies a larger area than phloem. The pith comprises hexagonal or orbicular parenchymatous cells with intercellular spaces (Fig. 3, Table 1). Leaf anatomy: Cross-sections of the lamina sections of adaxial and abaxial epidermis of V. globiferum have showed that both epidermis are covered with compound, glandular, and eglandular hairs and they consist of uniseriate rectangular or oval cells of cuticles. The vascular bundles are collateral. The curved vascular bundle is surrounded by parenchymal cells. The xylem 206 KILIÇ elements are arranged radially and form a single layer. The midrib is well developed. The parenchyma layer around the vascular bundle covers a large area. Cells of parenchymal tissue are polygonal and tightly arranged. Cells of the upper epidermis are clearly larger than the lower. Under the upper and lower epidermis is the hypodermis. The mesophyll tissue is divided into palisade and sponge parenchyma, with 3-5 layers below the upper epidermis and 1-2 layers of palisade parenchyma above the lower epidermis. Palisade parenchyma cells are elongated, cylindrical or quadrangular, and irregularly. Between the lower and upper palisade parenchyma cells are sponge parenchyma cells consisting of 2-3 rows of cells. Sponge parenchyma cells are oval or polygonal and there is more space between the cells. Also, idioblasts have observed in the mesophyll tissue of leaves (Fig. 4, Table 1). Fig. 4. Cross-section of the leaves of A: V. globiferum, B: V. lysiosepalum. G: Glandular hair, Ch: Compound hair, M: Mesophyll layer, ad: Adaxial surface, Co: Collenchyma, P: Parenchyma, X: Xylem, Ph: Phloem, ab: Abaxial surface, Ue: Upper epidermis, Le: Lower epidermis, Pp: Palisade parenchyma, Sp: Spongy parenchyma, Hp: Hypodermis, i: idioblast. Cross-sections of the lamina sections of adaxial and abaxial epidermis of V. lysiosepalum have showed that both epidermis are covered with compound, glandular, and eglandular hairs and they consist of uniseriate rectangular or oval cells of cuticles. The vascular bundles are collateral. The curved vascular bundle is surrounded by parenchymal cells. The xylem elements are arranged radially and form a single layer. The midrib is well developed. The parenchyma layer around the vascular bundle covers a large area. Cells of parenchymal tissue are polygonal and tightly arranged. Cells of the upper epidermis are clearly larger than the lower. Under the upper and lower epidermis is the hypodermis. The mesophyll tissue is divided into palisade and sponge parenchyma, with 2-3 layers below the upper epidermis and 1-2 layers of palisade parenchyma above the lower epidermis. Palisade parenchyma cells are elongated, cylindrical or quadrangular, and irregularly. Between the lower and upper palisade parenchyma cells are sponge parenchyma cells consisting of 2-3 rows of cells. Sponge parenchyma cells are oval or polygonal and there is more space between the cells. Also, idioblasts have observed in the mesophyll tissue of leaves (Fig. 4, Table 1). ANATOMY, POLLEN AND SEED MORPHOLOGY OF ENDEMIC SPECIES 207 Pollen morphology Verbascum globiferum Hub.-Mor.: The pollen shape was oblate-spheroidal (P/E: 0.95) with a polar axis of 12.88 μm and an equatorial axis of 13.53 μm. The aperture type of pollen was found as 87 % tricolporate and 13 % tricolpate. The colpus was in long-acute ended with a colpus length of 9.48 μm and width of 3.28 μm; porus length of 3.71 μm and porus width 3.51 μm. Exine thickness was 1.01 μm and intine thickness was 0.52 μm. Ornamentation was reticulate and reticulum was shallow (Fig. 5, Table 2). Verbascum lysiosepalum Hub.-Mor.: The pollen shape was oblate-spheroidal (P/E: 0.94) with a polar axis of 12.95 μm and an equatorial axis of 13.70 μm. The aperture type of pollen was found as 90 % tricolporate and 10 % tricolpate. The colpus was in long-acute ended with a colpus length of 10.27 μm and width of 3.25 μm; porus length of 4.07 μm and porus width 3.61 μm. Exine thickness was 1.09 μm and intine thickness was 0.61 μm. Ornamentation was reticulate and reticulum was distinct (Fig. 5, Table 2). Fig. 5. Scanning electron micrographs of pollen in genus Verbascum. V. globiferum (A- Equatorial view, B- Exine sculpturing), V. lysiosepalum (C- Equatorial view, D- Exine sculpturing). Seed micromorphology According to the measurements made, the dimensions Verbascum globiferum from 0.62 to 1.11 mm in length and 0.37 to 0.78 mm in width. The shape of the seeds in the genus Verbascum allows a distinction to be made between species and subspecies subtaxa. Prismatic-ovate, oblong, with ±shallow alveolate, multiple linear, deep and broad backs are the shapes of seeds. The seed truncated and obtuse beak. The seed is color brown. Because of the irregular, exserted polygonal, and small rectangular cells, with densely and distinct vesicles, a networklike appearance is seen and the seed surface coat is longitudinally alveolate. Inside the cells are 3-4 transverse lines (Fig. 6, Table 3). 208 KILIÇ Table 2. Pollen morphological characters in genus of Verbascum (min (mean) max). Species P (µ) E (µ) P/E ratio Sh Clg (µ) Clt (µ) Plg (µ) Plt (µ) Ex (µ) In (µ) Apt Or V. globiferum 11.56 (12.88) 14.35 12.18 (13.53) 14.83 0.95 Obs 7.35 (9.48) 11.20 2.08 (3.28) 4.56 2.23 (3.71) 5.57 1.87 (3.51) 5.25 0.80 (1.01) 1.58 0.26 (0.52) 0.82 87% Tr 13% T R V. lysiosepalum 11.58 (12.95) 14.75 11.78 (13.70) 15.76 0.94 Obs 8.59 (10.27) 11.89 2.04 (3.25) 4.12 2.91 (4.07) 5.55 2.40 (3.61) 5.74 0.84 (1.09) 1.66 0.35 (0.61) 0.98 90% Tr 10% T R R: Reticulate, T: Tricolpate, Tr: Tricolporate, Obs: Oblate-spheroidal, Sh: Shape. Table 3. Seed morphological characters in genus of Verbascum. Species Grou pa Length (mm) min (mean) max Width (mm) min (mean) max Colour Shape Seed surface V. globiferum K 0.62 (0.91) 1.11 0.37 (0.56) 0.78 Brown Prismatic-ovate, oblong, with ±shallow alveolate, multiple linear, deep and broad backs, often with truncated beaks, and some with obtuse beaks Irregular, exserted polygonal and small rectangular cells with densely and distinct vesicles. Inside the cells are 3- 4 transverse lines V. lysiosepalum K 0.70 (0.94) 1.20 0.31 (0.56) 0.68 Brown Prismatic, prismatic-ovate, oblong, with shallow alveolate, multiple linear, slightly deep and slightly broad backed, truncated beaks Irregular, exserted small rectangular cells with densely and distinct vesicles Table 4. Capsules morphological characters in genus of Verbascum. Species Groupa Length (mm) min (mean) max Width (mm) min (mean) max Color Shape Hair V. globiferum K 3.89 (5.43) 6.49 3.10 (4.26) 4.87 Brown Ovate, oblong Stellate, tomentose, branched, glandular V. lysiosepalum K 4.11 (5.44) 6.11 4.28 (5.47) 4.85 Brown Spherical, ovate Densely stellate, branched, glandular a According to Huber-Morath (1978) According to the measurements made, the dimensions V. lysiosepalum from 0.70 to 1.20 mm in length and 0.31 to 0.68 mm in width. The shape of the seeds in the genus Verbascum allows a distinction to be made between species and subspecies subtaxa. Prismatic, prismatic-ovate, oblong, with shallow alveolate, multiple linear, slightly deep and slightly broad backed are the shapes of seeds. The seed truncated beak. The seed is color brown. Because of the irregular, exserted small rectangular cells, with densely and distinct vesicles, a networklike appearance is seen and the seed surface coat is longitudinally alveolate (Fig. 6, Table 3). Capsule morphology According to the measurements made, the dimensions vary Verbascum globiferum from 3.89 to 6.49 mm in length and 3.10 to 4.87 mm in width and V. lysiosepalum from 4.11 to 6.11 mm in length and 4.28 to 5.47 mm in width. The shape of V. globiferum capsules is ovate, oblong, and the V. lysiosepalum is spherical and ovate. V. globiferum capsules is brown, while V. lysiosepalum ANATOMY, POLLEN AND SEED MORPHOLOGY OF ENDEMIC SPECIES 209 is dark brown. V. globiferum is covered with stellate, tomentose, branched, and glandular hairs, while V. lysiosepalum is covered with densely stellate, branched, and glandular hairs (Fig. 7, Table 4). Fig. 6. Scanning electron micrographs of seed in genus Verbascum. V. globiferum (A- General appearance, B- Surface ornamentation), V. lysiosepalum (C- General appearance, D- Surface ornamentation). Fig. 7. Stereo microscopy photographs of capsules of Verbascum globiferum (A) and V. lysiosepalum (B) species. Analyzes of anatomy, pollen, and seeds of the endemic Verbascum globiferum and V. lysiosepalum species studied for the first time in this study make them comparable to some of the other Verbascum members investigated. The present study indicated that V. globiferum and V. lysiosepalum have a very large xylem area at the root and a thick cuticle layer on the stem. In addition, the same features were also noted by researchers (Özdemir and Altan, 2007; Alan and Gökmen, 2015; Küçük, 2017; Tekin and Yılmaz, 2018; Aktas et al., 2020; Küçük et al., 2021). In some studies (Alan and Gökmen, 2015; Aktas et al., 2020), it was stated that the cambium was indeterminate in the vascular bundle in the root, and in this study, it was found that the root had similar characteristics for the species. In the leaf, cross-section analyses of the species, densely glandular, eglandular, and multicellular 210 KILIÇ branched hairs were observed on the epidermal cells. Similar results have been reported for other investigated Verbascum species (Alan and Gökmen, 2015; Küçük, 2017; Aktas et al., 2020). Idioblasts seen in the mesophyll tissue of the leaf can be considered a factor for identification within the genus Verbascum (Lersten and Curtis, 2001). In this study, the presence of idioblasts in leaf mesophyll was determined and this feature was also noted in other studies (Kheiri et al., 2009; Yılmaz and Dane, 2011; Tekin and Yılmaz, 2018). The pollen of endemic V. globiferum and V. lysiosepalum species are isopolar and radially symmetric, oblate-spheroidal, tricolporate (there is tricolpate), the exine exhibits a tectate structure and reticulate ornamentation. Baser (2021), in his study on Verbascum (8 species), established the pollen grains as tricolporate and tricolpate. Aktas et al. (2020) determined the pollen grain as tricolpate in their study on the endemic Verbascum species. Aktas (2019) noted the pollen grain as tricolpate in his study on Verbascum species. Özturk et al. (2018) reported that there were tricolporate and tricolpate aperture types in their pollen study on V. pycnostachyum (K groups). Al-Hadeethy et al. (2014) stated that the aperture types of Verbascum species (20 species) were tricolporate. Aperture type of Verbascum reported to have tricolporate aperture type (Kheiri et al., 2006; Asmat et al., 2011). The pollen of the examined species were determined as oblate-spheroidal shape. However, Aktas et al. (2020) recorded the prolate pollen shape in the endemic Verbascum species examined. Nevertheless, Baser (2021) recorded the prolate pollen shape in the Verbascum (8 species) taxa in the studied. Öztürk et al. (2018) stated that pollen shape was oblate-spheroidal on V. pycnostachyum (K groups). In addition, Al-Hadeethy et al. (2014) observed the presence of prolate-spheroidal and oblate-spheroidal pollen in Verbascum (20 species). Morphological pollen traits observed through LM proved not very important in their taxonomic use, but sculpting examined by SEM was found to be more significant in the classification of taxa (Pehlivan et al., 2008; Baser, 2021). The overlapping of exine thickness was found among the taxa studied, thus this character was of little taxonomic value (Al-Hadeethy et al., 2014). Results obtained from the present study agree with previous studies on some species of Verbascum (Asmat et al., 2011; Al-Hadeethy et al., 2014; Öztürk et al., 2018; Aktas et al., 2020; Baser, 2021). However, the studied species showed a neat reticulate exine sculpture pattern. But V. globiferum reticulum was shallow and V. lysiosepalum reticulum was distinct. The seeds were brown when mature. The size of the seeds usually ranged between 0.62 to 1.20 mm in length and 0.31 to 0.78 mm in wide. The measurements taken in the present study were compatible with those of the common species in the studies of Attar et al. (2007) and Kheiri et al. (2009). Yet, the measurements taken in this study were larger than the mean measurements of the species common in Cabi et al. (2011). Seeds in shape from prismatic-ovate and oblong in the species studied, and they ended in an truncated, obtuse beak. The seed coat was longitudinally alveolate. Usually, most seeds of the species are prismatic-ovate (Table 3). The result regarding seed shape is more or less consistent with the results of Attar et al. (2007), Kheiri et al. (2009), Cabi et al. (2011) and Baser (2021). The capsule of V. globiferum is covered with stellate, tomentose, branched, and glandular hairs, while V. lysiosepalum is covered with densely stellate, branched, and glandular hairs. In some Verbascum species distributed in Iran (Attar et al., 2007), the capsules have a similar hair indumentum. Based on the available findings, one of the reliable characteristics for grouping in Verbascum is the indumentum of the capsule. 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